SF 768 
.2 

.C3 B57 
1923 































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































D.:?iLGFor ih riVi DULTS ahd c*:>als 


i *, 'i :.n 


C - 


?, i : d :. ,:• tr ;- , L . 






A LIS dKTATZOS 

I ub ittod to the Faculty 
of the 0£d<m Graduate chool o* Science 
in Candid&cy for the 4>ogvoo of 
Doctor o ' Philosophy# 


fJKPUH £<T Of- ZOOLOGY 





rhor:uj buue -iaaonpotte 




J .4 J ■ si 


Xf£3 




i?m-b , 

©p* ? 96 j. 






The Cevclopr-ent of the *■^productive Canals 


X 



in the F oe-n^rtin with corsporiooc of iho Scroll• 




i-ofi iti n of Fre e-oar tin 


Older Views of its Mature 
’lature of the abnormality 

Lillie's ihoory of the Preo-mrtin ami evidence for It 
Previous fork on Jro, enitul Sy»te of Cattle Embryos 
Purpose of this Study and its lap: rt nee to the larger 
Problem of Sex-differentiation 
Central Conclusions from t is Study 
ii-iTdilAL iCii> l&Tl'iW'S 

Source of Material 

Treataeot of Material 

Moa; uresonto of dsbryos 

Reconstruction Drawings• flow judo 

Tabie of AriBuls Figured <*nd Stuaied for this Paper 


Table 2• 



■iOnadc of Feesale, rule a d Tunica a.bug! ea. 

Secondary sex-cords, etc* 



in fesoie, ’ale a d Froo-nciHLin 
^lioriaa Uicta ana ..erivativos; tubes, uterus and 


Vagina 

Vac ua vs iMslis, Hound tiganont, a i uteer aculur. 


t’ate o-" the deao sphroa 






































1 





































JrcgonitAl strum, r-ulbo-v©.-: titular, ulbo-urothral, and 


Pro: tate glands and futurathral i Iverticaiu^, :upra 
urethral divert!culun 

Phallus and rtornal nitalift as far an made out. 

::u tt/.f ^ U DIXJC.. 10.u 
















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: omcrioj. 


In cattle a f »ir ile born co-twin to n sale i sterlie in the 
rr-ajorlty of caeots und it? known 'iron,-, cattlo'os an a *froe-martin*• 
By ore authors all females of two-aoxed twins in cattle aro called 
frsa-mrtins* The proportion of the cases in t? ich uucb female io 
noraal is iven as follows; us&n (*44) one i • oi ? jht, L&er ( f 13) 
six in ninety-one, Lillie (*k 3) six io thirty-nine. These with one 
an thirty-six as fcr.;nd by the author* rake a total of 20 in CS7, or 
about 1 in 14. rtj-i roascn for the atari. ity is quite conclusively 
3uo»a by Liliio to be duo to definite and sufficient 

v .scular u:r» •• torbetween the circulations of the txi t in the 
placenta! eerbranee* in the absence of which tbs foraale io noraal. 
ihe -ale i • always normal. 

By toe claer autr.ors the froe-martin was classed a - 
b .rnauhroaite, or posude-her* aphrodite * as r-o ified r«J.e 
( .uTvan, '<.4)(J. Barry t art, ’10)(Bateson, # 13)* or as modified 
fecale (dunan, *4). Th ? last nursed author classed the* - m naies 
or ee females, according as the orpins are ore or lass ferule 
or «;ile in type. S'-degolborg (*6l) classed the ac transverse 
h sr aphrodites • 

atrxin (*4i) deacribthe case of a . uppoxed froe-r&rtin 
» ict hs obtained fre.m a i >ul©r when a year eld. » ich had an 
apparently feaalo sot of external ^onitaiia. revs, ventrally 
alo«u the body-wall, with ar acceu ory urethral openin dorsal 
to it. !1;e internal urogenital organs »sre decidedly ole, but 
o evide ce can be ^ot fro® his figure ae to the d.-velo wont of 






an uidor or scrctua, and tne latter would fee a strong criterion of 
alanesa. Roller no londler (*3 5) state incidentally that they 
have note; codifications of the external genital! in tfc nalo 
direction, but they do not caccribo there. A c* & is described 
and interpreted as abuorml rale by Pa.nucBon (*10) of a foetus 
twin to a nerval bull calf, which hid no prance, aero tire, vulva, 
vagina, developed cervix, fallopian tabes or hydratid, id no 
prostates, but which had a hert hidden onic in the urethra, on 
udder ana teats, a uterus with horns and holy, spinal vesicles 
viui ieforeos# epididymis, an. partially descended gonad* fith 
these few possible exception*, the ferule Is aodifisd in a re- 
xrk&bly definite and lisitiKi manner, so that her internal genital 
organs are cede over in the scale direction, on her external ones 
rsaain distinctively ferule. 

as to the cuu o of this resorbable situation, Lillie (*lt,Vl7), 
in America, and Keller and Tend1 st ( 1 3 ) in iuropo, ^uitc iado- 
penior.tly, <Arrive.; at the s&r-e conclusion that tho fro<.-martin 
ccoditi-n is due to the action of a aul© boraoro, free the ale 
co-twin, passin. by vascular an^stoaoei o' the placental vaeaelr, 
to the fo ale, which causes -odification ofthe internal go.orutivo 
or : uns of such fo als in the r direction, without in any way 
interfering with the norm! development of too t?alo co-twin. 'taller 
arid Tangier d rot *y where they think t> e hersone is produced 
nor nr.sr it begic.s t act* hix:io (*1?) postulates that since tne 
interstitial colic of the ~aa*ral testis develop lor. bo for thone 
of the- ovary, the, re protahly the vat o'* ale nortaono production* 
Lillio arri Ha co (* f) and Baseon (*2 ) chow that the interstitial 
calls of norml -.tie cattle c'bryos appear bet»*e • ?*’ f att 3*0 ca, 

stators, wh eb i about the tisw of the initiation of external rexaal 



3 


4iffererrtiation, while they do not appear in females till } cr M 
•tiich i. n r ti e t «-■> of birth* Lillie (• f * ) alienr. ti at toe horacne 
is already effective in a free- -riin or S.?5 ca«# and pcinto out that 
sy.sOtic ftos-dctaminlng factor:} aro ol«© ir. a e rtain degree aex- 
differentiating factors in ssasssals as In .resets, and that, nhlle 
in th ala these factors ere irol I - reinf • 

hormone action till abo t the tiro of birth at aarll rt, is *ho sals 
they are reinforced early# e seers to thin that th© hc^no ray 
bo introduce in the care of the free-martin before ©axual differ - 
tiation, but trie can hardly be true for the appearance °* * n t- r ~ 
atiti&l cells i-s in Itself a sexual differentiation, an: ir t o 
froo-' irtin, while tna ar.-.t ;is iocs t, fce pUc as early as the 
1.5 c.3# sta...e, Lillie (* )• ar.d pr bably as early, at* 1.0 c~ ., 

confirr-ea in t o writer* cello, tics# th hornene cannot be intro- 
uucad before the appoara.ee of the colls that produce it. In the 
free-cartin while the effects aay appear early in esse car *c and 
are the case effect.- as produced in t r* sale, in other cases, 
they dc aot appear till the ferric ha» under.. ore s coca id arable 
aroorst of sexual differentiation* Prscir ubiy t o hor~o- o has 
net been introduces till near th .t stars• 

To date so exact and ninuto a.-.to: ic>l ccn c isons of the 
our l i. r stages' of normal ala and fe^sele o bryonlo and foetal 
^.evelo parent hare been rsade with reference to the 1 fferertiatien 
of ti,e sex-ducts and oro*.enit«l canals. T iu, wifi: the sr*~e .sort 
of >tu:ly of tho r: -rurtin. in nectar.ary* o 4 ” cc- m as a W:ir 
for tho oor ‘ieto of toe course of the freo-cartln con¬ 

dition iind Its variation. 



4 . 


cer&aefeick: r (*1 ) ».ork@d out th© development of ti» foaalo 
Wolffian astc Ufillerian aucts yusa > ©nitfcl ppurutus i:. cattle and 
do cribo the changes found in tne norxsal feasaie. biilia {’17, 

) took up too protti an to-'leal odi loot! n« of toe fro©- 
sartio te compared : tr. »c;n© sales and fossalec on a baaii for hie 
theory of tie free- • .rtis# and a riesut ittt y O’* a .75 05 • iYee- 
•i-ortin a* n»i ^ stoma:. fhitalia. fh^.-in *17) inci- ton i her 

pi^r o- tho footal frie-eartin * onad sa..^ notes on tr.e ducts 
adjacent to the ^onad• Ail oar . ier cttiuies otn-iot for tr© coat 
part of Ari^to^ieai description* o individual free-martini' without 
. an., attarpt to correlate t^a situation with -orr,gi odes of 
development. 

-rcas d nsection cannot show the real state cf tho duets and 
their derivatives and definite knowled e of these c -rd lions at 
o ccsssiro nta, os in both normals* and frec-oartia## a d a com¬ 
parison of one vita a other at equivalent stages. in necessary os 
the m:sic for dc sire conclusion© on to tn© course and caucus 
cf trio dificatioiu 

At the suggestion cf Professor r* h. Lilli©. the writer 
started to investigate the norm . velcp-ont of the cex-dmots 
and ;-,er:itai canals ip sale o*:-d fere © c#lf-ombryoa # up to a eta ; o 
when tho or,.-os or© well started t war:: their efinitive co- - 

dition. r or top rgoso of csr arin* the odes of development in 
the two coxes. At t e collection of n t rial for the study out- 

Un-d at eve went on. free- rtin w r© taken which fitted into the 

V 

incorporate, into the study. In one can , the cor pari son cf two 

ad to uste » and cor¬ 
relate t;,o variation fouid in t,.o ;© of* tac eaoo else* 






















































































Tho writ r wishes to otpryus his gratitude to Prof ms cr Lillie 
for his suggestion ef the ori*:ir-aI arotlc*c and f, or unfailir urd 
fcindly advice and instruction ibroufhout the work. 

f tar'ing with th premises as a working hypothesis, that th© 
frse-aurtin sa u ay otic.illy .letera!red f.-?aale firrt, and that 
she wo* Id have h A the normal female mode o* development throughout, 
but for the action of the mal© ho-rene injoctoei into her in uiero 
by «uy of the placental vascular ar^ntemooie, to investigation has 
boon an *tton t to find out the offset of thir calc ho me no on tho 
fer.le ;,ubryo* ic dovolc .rent. This is done by 'tie pii.iv to an- lyoc 
the 5eri©a of natural experiments prei«erteJ, with reference to the 
factors influencing th varying results, and different d,>.rees of 
the oa'se result, proiucad, using tho normals u* control;;. An attest 
has tnereforo been made to test tr.o worfcin/ hypothesis and to answer 
the following quo .-tirns in -addition;- 

(1) 'that are the no ml fersule a- i male soles o~ dcvolopnentV 
{£) fhat chanfc© from the expected normal female ode have 
rosultodt 

te rurlatioafl in the ol b 
effects of the herootj© in the froo-martins st .iicit 
(4) Is tiio action of the hormone limited to any seta of 

structures? If so, to what o re©, ni in what manner, 
does it affoe . each act of crguno? 

Ti.ifi investigation leads to tr.o fellow in general c^nc ion ices 
in addition to producing evidence that tho working hypothesis sac 

4 

so nd;- 

(i) Tho crtsal for»lo development of the yftl orion -uct- 

: , - : ry »©x~ cords U 1 

, 

the develop ©nt of th- female type of ainu© uro *e ii&li a, with 
suburethral -div rticuluc, vestibule, vulva, bu 1 bo •vest; fcu iar laris, 
and clitoris: and the development of a roun Ugu c- t; the de- 






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•i 1 

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6 


generation of the Wolffian duct erivativoc; and the ear~v lose 

to 

foiffian duct, would have gnus on apparently n.t'P the c r.trc-i of 
*7* otic factor , unaid d by ro 1 r:frrcec- nt by se'Jt-bcr:n©ne action, 
at least till ns*&r to tins of birth. 

iha Qomal sale nod© consists of the dev©lop-ant of the 
o If flan duct derivatives, the early eparatior. of the so coords 
fr p the opitheliu of the (©nad, and failure c“ soeoBd&ry sex- 
cords to fws; tbs early u exoneration of the Ullorl&n duct 
derivatives; the development of tr rtale type- of urogenital sinus 
derivatives, without ^uburethr*! iiv.rticuiu , and *itn short urethra 
proper, and Ion,, pro:;tatio and cavernous urethrae, si-noi flexure, 
bulfco-urethraI gl ads, ©:sd Jistinotive little ouch at the bend cf 
the urethra; the ventral igraticn of the penis ana growth of the 
perineum; the develop- rh of scrota and di tinctlvo fT ufc ornacutas 
and aaocutt vaginalis; und r the control o .y. otlc factor 1 roinfbroea 
curly by vale hor^o;© action. 

(2) The changes flrce the expected feral© coae are; failure 
to appear, or degeneration, of the «ooo dar cox-cords in it gotudj 
. 

shoeing ai*. *\u of having developed a in f& aies to varying iegreoc 
before d e crating: the cevslopns- t of the • olffion duct derivatives 
as in aalec, with signs of early partial h@|eaeration aa a ferules; 
practically no chan # in the fetaslo rede of development of the uro¬ 
genital sinus derivatives: and b velopmcnt cf a tsore or I^os ''.ale type 
of gufeer; tculur and caccus vaginalis. All #; Ofi t e effects of the 
sale hersone parallel to t ; -ose in the ale. 

(i) fro© the reruiti brought out by t Is study it U believed 
toatthe factors underlying the variation in degree of effect cf 












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7 




this horaono in different fr**-mrtiii« of Ur- sarsa else and In t! oj© 
of different iiswo arc;- (a) Tits : •* initiation of her? r.a xiion# 
which .n turn (i«j ods on ti :» of hormone production • r» the r:l« eo- 
twin# 7p on ti~e of cor:emotion of phe rt-*l va«cuior sm- loses!c. 

(b) The leni t of 1 :»G tba hormone b*ts$ bean acting, just os in the 
r *c r^al c.,Xe, -hero tine is* t factor in the state cf the ducte, otc.* 
at an/ sta^e» but with a v&riublo condition of too organa upon e iofc 
it has to begin action. 

(4 } In all cases oh.-: rvod. ti e action of this hormone on the 
organ© ( >roductio 1 k lied to orgfr: t MMOi* tail origin* 

aitd fails to affect, those of ontoderoal origin, except, perhaps, to 
cause lightly earlier appe ranee of para~urethr.il ducts• * ich, 
owever# arc net sale in type, tout definitely fe ai», xtornel ;erl- 
kla«_ iu.al imi .-.i rvod, are uoaff Ot i stale d I 

though it if: not lost right cf, that son© such nodifie&tic- 5 ero 
described by ot; r cbe rvorc» it is b©l I ovoo: that they *&ay well be 
of sy otic ori. it? ar.i not due to nale hornone ixetior in uterine lif- • 
uch 'rrcdificaticn; ac • ear in sir: le-bom individuals, art are there¬ 
fore to bo expected in about the sane properties 5; sujeng two-sc re 
twins, whether wit cor&one action or without. 

The u; parent p^oificity in tho her-nene action is due to 
specific reactions of *-he ti-;s‘ M affected, in -but eoae arc 
sticsilat^d to incr :a e.j oveXopaent and differentiation by it, 
while others are inhibited cr-» cause : tc de.en-rate# and still 
others apj^esir unaffected. The chn--. v or; in or. ral body slap note 
la free-a&rtisa# and in equivalent organs in th 5 two sexes, in w .ich 
the fr o-®artina ro.-.e • ble c.’sles aonsatut, *©uld show that other parts 
beside sox—organs arc affected differentially. If th or ^ are part© 
unaffected# they nay ell fc cy otically controlled or past a critical 






































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1 is Tfouli ai'O iccount for 


ot-i •* fcv-fv.re the hormone richer ^er , 
the f c* tuat cc'" piste flex-itivarsiOr. is cl "ound in free- r&rtine. 

The fact that 160 • fleeted cr,/A. **- in free-vurtins lie anterior 
to tna v 2 Gt cula* vjiil account for their bo in* clssaiM* by early 
writers o. transverse fcarr. .ohrodits 3 , 

1 is tstudy uli-O wok t. at, net or iy 3 the r,I« acr^n-e in¬ 
f-tit the develop *?nt of .Sllorian duct d rl7a 4 iv;s ar.d ovarian 
, 

. , : it causes ictiv . ••rition of 1 -as 

structures wfcen ti nv arc .lready forr.-ov in t.. e fornix embryo. « is 
would land seaport to the idea of rteinacis (’if and later) 01 
Lillie (*•• ) in at tno tc.?tis, tfcrou-, n it s horse •% ex tu an action 
that is uiita onistic to the ovary, at lea t in sarly embryonic life, 

) ©ore (* 1 ' ■ 

lievir.i that* in all c .g > 3 , the sale borcoao iu atisylstirr in effect 
and never iu ifciti ng, «ot only is It inhibit in. in motion, but even 
actively ante,, onistic to tno scondsry eex-cor^s .Ires y d veiowed 
to a considerable de.r e i an embryonic ovary, 

Thor® is no evidence found in the pursuance of Inis' study of the 
formation in cattle reciprocals of tno frss-.-ertin *» inferred 
by ..artean {*<0) for the Qpposu? a 4 wide; tly sooepte by bin as 
possible in an and 


:4AT. hlAi. *,1i> *T OL f. 


All tbs orbryoa upon union iriia niu y is based wvre col. acted 
by tno writer or, t.-v teef- floor of -if* a .a cnapony. iv 

wnicu t o, between April ,-1 and nsril, If: 3 , t e writer is in- 
defctei t. i ,e-, ard to Ja~e falker* their foreman on t; o killing 
floor, for - ind oof.-erutioa an- valuable a30i.5tr.nc9. 


.. t ri ware ©; ene J as 3©'n as take free! t e co^, 
inutsc after .isr throat h-.d boor cut, t * chorions t 

Oi O'? o , ar .1 t -j embryos Is n than . 1 i~ 4 w 1 

, . * re, ' • ' ' ' . 

ana fixed. 


about tO-^6 
afeen out and 

1 n»t;., killed 

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anal length ;,ad the ubdorroi* evened* t> o Itv r and intestines taken 
out. and vers then iaseii ately pat intc Bouin's flail# Larger 
si pea hud U,a front quarters cut off .t t « ucfcilicua l.rv.»i t the 
syrphysis pubis ana iachii cut throu* fc* oral r rafxd, to lot in the 
fluid# a.- ; a * then put in couin'a fluid# 1 in insure? k lling 
una fixing fr«:.h * for. in t o;..a a. enhryo -< of a-d c '• h - '» the . 
heart continue to beat far fr.r. 12-26 ninutes ftor their thoraxes 
«ere or jn«-h In nc care un Material# that wan kept are than tsro 
hours b fore , oin. into' the fetin'a fluid# b en, used i this study# 
After "4-26 hours In bouin*a flu d kept strong * y chan. es into 
nee fluid* tb ; satoriul waa mashed la 60' an, ?0f alcohol and. then 
in *3Of alcohol to which & trace of Lithiur. carbonate he,, bean 
a ;ed# for two aeys to reoove Picric acid. It *u the it apt in 
80' alcohol till discectoi for dreeing or photographing before 

t 

oteisiRt; -rid sectioning# After draRin?,, fer rrcoru of ^resa 
ar.utosy# th ; larger -atarial, 7.6 c . end over# bad tba right .onad 

f the semt&X oords* eat “ 

be-id 3' irately cii •. -c • :: tr,-. ....... ro '’.a: r.e.j 

in bulk in Alua Cochineal f or ’ 4jA20 hears, then dehydrated in 
alcohols *nl yr.tr-stic oil of inter green an; asr -ridin soft 
paraffin for s rial- ectioning; vross 16-^0 -icra thieve# esc act in 
cases shore the otter aide *& sect! ;.sjd 4 inner for scaparison# 
Aft:r bou ati» the section erriea were counter., t^inea ir Orange .., 
cr -JLcfrtgrhn, but t.* « 1 dfr stain »a at «;.d- n.-j , a it i • not 
..ive ar clear d5 fforentiatir* as the ;range L# 

•tec sure-- ents a ivon in 'his study are ^errh sal-anal lengths, 
for those that bax straightened out# and fr a tb* c rvical flexure 
to th e oppo ita curve 0“ the ru«p, in seller once# These ”■♦cures 
ware ocsiest to are accurately .it the tir>e of collection of tine 
killin.*-♦’loor, i- the car.es of the larger one , an: r or the smaller 
ones# rhorc .r.t c old not he accurate till V. 11»J and fixed* 



1 


k*0*2iXB'> u- ‘i ed tis?mca *©r© too nearly tranapareot, it man ire vest 
constant 1 r.gth -wt y tc .uxsrz vocurataly, If vary yo-rn/ t* ©a 
tc • .’ rtu -«ied sc-' © jtdjut t -nts will bst © 005 ;*:ary* 

In all caoen axooft those of ‘he 2b.O cr* fencl© (fig. 9) and 
• * • lift act Of I •'*. | to 

beta sid-** son .-©asn'rca for *j. e reconstruct!'ns, hut in the so cases 
♦be rii.nt olio wa * substituted h©c ; it* s of ^cci seat to tr« .• loft ..ions*! 
ar.u genital oore*. rhis mukes th-rs© rsc©7*e 4 ructi loo or* because 
the right oide has a loss a vane id sta e of retreat of the organs, 
t-iiati.sr *its its other evi nonces of slower dweioperni* fheao 
facta are tatsa in connection with th© affected parts in the d©e- 
orlption o** the findings* 

l n ,j ill ; tr.it ons for tevi* pa|>er are ©do fc • 'r* hen r i too a 
• i * urea 1 and- 3 fror. the action-;» and t © rest froe ft-- orifinal 

crawln» to scale by the author. 

In n iking the original drawls s, f> or larger opeci-©nr than 
d.l co. # the rector *• - taboo a ref rer.ee lift.. , fr*.» vi ich to 
a©**a .re the dorso-ventral displace". nt in icra* as shown by an 
ocular ssicroa .tor calibrate * a f ainst Zeiss :» 4 i. r-- ©..-er* 
ouch ;ivision of '&<• ocular icror.ot -r ccrresv.o-uliRf tc ho.? 

icrs 4 . Length along tt© antero-poct rlor aicia was obtained by 
court in... the z rial ..actio .s fros the po tori or tip of tfe" Phallus 
forward, ana swltipiying by Ina thickness in vicra per motion, 
ura^ings from ?*t on. erveeimma up, war a ado to a scale or fi*« 
ir tot., dimensions, with the exception of the lo* cm. rale* 
which, with tb- smaller s : >©cl--ena* sac vmdfe to a .ucule o; ten* 

. . ; ;. t-> r •: : ■ f I - i* ; c 4 - 

Ca '0 of the 53.0 cau feaale, where the olffiasi duct derivatives 
atro placed vertr >1 to the vagina *bil*> they are really in the 
vontro-lateral walls of that organ. In the ca a of the guber- 



naoulun th re is aone corventi- n umd aa *.© the a ocus v-vi nails 
and %ho li: assent, 4 o show the cavity in the ,'uccug curro'cndin^ 
the r uberrmcultm ligAK-.? t. Tn nuch rec-n trueti n it w& im¬ 
possible to sfho* ho* they joined the cord in either "aloe or 
females* In all c sas the also relatio .s o r *r. age structures 
are to tne sa:' : e scale, a© distortions dm to faulty p ition 
in rea^esits ar net corrected* In cacaa rners a part c a struc¬ 
ture i« ;is«octed e«uy or injures by accident that structure is 

£ ? 

I . 0 - fe ale and 20# j co* .ale, represent ...ifror of the 

org.-Mn, an to lat r *1 icplac.:aent, so that th organa f • he left 
side appear beyond tne iucto whereas they are rc-liy nearer and 
would cover ther* the duct) are represente: a cut by -ajia 
vertical Ion itudin.*l section t? rod... h ti.vir tire length, but 
in oases ^fcare the lu ?n leas in 4iabater than the true’cnees 
of a lin t th: c 1 a- o ' th ■ , ri r issai drawing the duct ap r w?3 
solid. Co the drawings aact not be taken a n ofc.-.v lute guide a to 
pr seme of absence o-' lessen. 

In ttio ca u o“ edbryoe lass than 5*1 c . forehead** al length 
the notochord waa v oi m raf rs c line, the anterior pot of 
-ach irawin , V^cause t a rectum sake a ventral loop near the 
p. f terior and of the somaepbro. • Cooo ;-li fc ht ui-tort .r.n ay be 
to twi tin of ecti a a ittlo in counting aiv.s iu: t i t&isan 
into con. ia ration in a critic . , sx-ainutior. of the 












■ 

’ 








a 


„ 


. 




* 









' 

* 













- 





T i-.Jw. 




\ 


mm 

List 

of 

■•ifeur-C ~nh Anio^l; for 3* 

udy. 

Fife. lio. 

Longtfc of 

o'nbryo 


tide 
u ad 


tnict T 3 s: ofinn 

1 

?.0 C2. 

r enale 

Loft 


.ectjtor 4 • az an. 

S 

9.0 

« 

re.;-” artm 

« 


0 

• 

w 

e.s 

« 

Faaale 

» 


i.econatructi r 

4 

3.0 

« 

0 

0 


0 

6 

3.' 

* 

0 

0 


0 

6 

5.1 

. 

8 

* 


9 

y 

** S 

4 


0 

0 


0 

6 

9.0 

0 

• 

0 


« 

9 

16.0 

0 

* 

s if.Lt 


« 

10 

18.5 

0 

• 

Left 


• 

11 

" 0.0 

0 

• 9 

0 


* 

1 ? * 

2.6 

0 

aid 

« 


tt 

13 

3.0 

0 

8 

. 


0 

14 

3.3 

« 

0 

0 


0 

15 

5.1 

0 

8 

0 


0 

16 

7.5 

. 

8 

« 


0 

IT 

10.0 

• 

« 

« 


0 

18 

17.0 

a 

* 

8 


8 

:9 

80.0 

0 

9 

;• ight 


0 

20 

0.0 

0 

Froa-n,ari\r. 

..aft 


S 

n 

9.0 

0 

* - 8 

» 


« 


11.75 

0 

0 

0 


8 

«s 

37.3 

0 

0 

0 


0 

84 

50.0 

0 

8 

0 


0 


Aniaala 
0*3 c. . 

3.3 * 

1.4 

1.6 * 

i ■» * 

> . 

c «t * 

studied, bwt not Figured 

Indiffira-it noth sidoa 

« 0 0 

, . * • 

» 0 * 

. • * 

0 • * 


f* 4 

• 

If 

0 

* 

0 


- 

*> » 

« 1 

** *? • 

• 

15.0 

••9.0 

9B 

6 

«r 

« 

ale(prei’Ti aa V 
* (cartalal !t 
" Loft 

Free-martin G 

8 

« 

i.r>ato'.y 


;13C ts 



















































































• 
































•M' 











;.;.YO£rv ,;i OF {feurtK aSj SHHiS. 


The onad. 

-Inca th© gonad of the Froa-jsartia baa been stadia' by H.apin ('1?) 
and illisr (* 1), and is not the smin ic ic of t is ©tody, it ie 
Introduced bora ssainly as a ©bock <•• the other work. It it partly 
ccnfirr itory of previous rotrslt© cu-d .a addition intr duoor, sore near 
fsateriai. 

In the indifferent eta ea;- at 3.9 cs. there is no oi r« of the 
ge ital rid^a, at 1. c . the .marital riug » just st-rt to thick-on 
on the ©©dial aspect ■ f the r-.wtwrpfcroc. They ar ;ui H vorkei at 

. . . --■ ■ . tro- r .i •;.: a 

neso.~epr.ro3 conr.acte. tc it by losexstoriaa. :<o sexual cii fferencea 
are visible. 

In f©aalea; - At 2.5 csj. bfco sale -n f oral a onads can be 
d is ting*'irked. he overy ha.- a n^rro* Incica &lba&in©& .. op».;rating 

too prinury e ox-cord-: *> » t; » *• *r*:.. nal •ipiifc’sliir? •• td :b i« several 
layers thick at this ota- »• fro© this ata^e nprnrh the ©• can ba 
di ;tia ; vj.- h .*! by the on-#i aactions .ter a &au o 'actitre. 

. x 

osil-lm/ >r# thick, th priaary iuaiea alba. Ut»& as ds-aeribe ; by 

♦ 

uh*pin (*!?) an. tiller (**!}, i a sere evident, 

5.1 c.• f© -alea have the * nrainal epitheliu prol furating in- 
wards arid producing th ra ..ic. o i ‘ seec-rsdary -cords or cor is of 
Pfizer. ? icb are cut off fro© the roll© of the ovary by th© 
loss intensely etalnon primary tunica .Hu., inea. ha medulla oon- 

e 























































































14 


The ©varies 7*5 cr* ** bryo. ! : & - a decidedly fesaie in type with 
a cortical l-.y*r eonsi tie* of covers! layer# ger Inal opithsiiw* 
and secondary se.'.-cords* ontsid? and separated by the pr stury ttmica 
alfcu f inoa frcsa a alalia of primary ees-cords and rote* T is tunic 
is be•‘sing vascular* The gonad id of the typo described by Chapin 
(*17) for the Roroal ovary of this filsed ©rbryo. 

At ?*0 c . (fig* 1)* the £#• inal epilfceliu i ntill connect'-d 

/ 

to the t iclear if yer . oserdary .ex-o©rds* the rivalry tunica al- 
ta&^n&a s Rare vascular, the ed -11a loosely p^c'.ed* ani tne rote 
rs a r.s near th o^ssertary* 

At 16*0 c • * the ovary is larger* but shoes the ssrse i -erol 
topography ir, a metier a - th- • cr. nc»noul fo^ie in Chapin* ( ' 1" > 
paper* except that the definitive ov\*.ri.n tunica alhu Inca ir not 
yet forcing* 

At 20*0 OR* the germinal epithelium is otll connected to the 
secondary «ox-cords which are thicken* tr-v primary tunica al¬ 
ba inesi is thicker and more vascular* The rote is near the 
mesentery in position* 

Particulars a to the latar stance are fivon by Chapin C*17) add 
are beyond the scope of this paper* 

In mlos;- 

Tfstes are first dictingtr sbcble fr a overien at bout 1.5 cm* 
by the fact that the germinal et iheliu become but one layer 
*;•• ic a ... ij.ii cell ; ’. : . it* ‘"ur c-1 to 1 ' * ;r 

eta ni g Iyer* the tunica albuginea, rsc losing a denser staining 
edulla* of cello which *1U later . ive rise to the primary *y-cords 
a j finally to the seminiferous tubules* The rote is more Cc trally 
placed than in the female, but Is hardly to bu i tio>»uieh 0 d from 
the surroundin cells at tbl: uta,;;e. 

































































































, 


















lb. 


«t T. 025* the cutting off of the garni al o itheliur frosi the 
sexrcorda is *o- • mrked. The t aloe &.r K arid or . 

i-sa darkly thaws in the female. The rote la nearer the center of 
tne i.onao and is etore clearly dint In uishatle frr the ties-cord 
rollers o r ‘ tr-j redulla than in the feral®. 

At 1 *8 cc. the serous epithelium of he gonad is one cell thick 
ard its cells yre flattened* The tunica albuginea is cronies out 
t-j a thin layer n -ar the periphery of the gonad by the larger sex- 
cord region* The rote i ; c.'itral and darker sia no tfcar in the 
feral e* where al.ro *fc© tunic is not so well -a ef red * 

At : *1 cr* the sex-cor -j are growing outward and fern a 
definite layer surrounded by tunica &lbu ir>ea. "be oex-eords are 
or anising :nt tubules neat tne t nica. ho eucr. definite arrnnfc- 
fsent ic evident in the fe as* 

At f*| c * the tunioa ifcu, inoa of the teati-r hue U-cn e • r 
vascular in its inner part n.-.xt the sex-cord* and be, nr tc fore 
the t nica vnseulooi, isaking tbs ole tunic appear thicker* The 
rete is ce trs.1* 

At 10. os* increased sits is tr.e only a arent ch-. ye* 

At 17*0 cr* tnere are def-nite tubu.ee extending fr ~ tee 
tunica rmeculosa t- the rote. These have primordial gerr cells, 
each sit. a large nucleus a rroundeci by a clear » no. This gonad 
re sorb lea Tnapi *s fi re of toe testis of a 34.5 c * nor ai, and 
Util * a 20.0 © . the one here it* U 

The fr e e--art in .onad• 

Lillie (* 13) describes the gonad of a .75 cw. frea-saartin and 
Oha r in (’1?) on of 7 a C . ana Vo reader is referred to their 
figures for t: ecu at* e»* 

In 9 * cn. free---artin do. 1 ( i,,. )* t. © gonad ie but 










- 






































































» 

" 


































■ 

































16 


slightly e \aller tba thut of the normal fossa•e, which it euenkle? 
ir, may respects an to erces-iection. .owever the secondary sex- 
cords, cr errs.' of Pflfjger, are already o t o f ff the ^or rnai 
epithelium by a secondary tunica altut.ir.ea of -:.«ie t pe, n j 4 ks 
r ir.ai a i the liar- has hoo---.;e flatte ed, Tri ' tunica i ipproxi- 
etoly like that cf « male of 3,£ c.r.* ana no tunica v sculoisa is 
evident. Ike seco -dary seat-core ssono is .j! :o.*t *s thick a in the 
normal fei. ale, bat in not a tch invaded by bio i-ves3«ls from the 
fOBCalsr ;; • y 4 ’> Uu. c lb 

outer ar. in cr heavily pi, cried, and appear to be under eing 
do^enerati n i a co tr petal direction, 'n id® this zc a ir the 
vascular primary tunica albuginea like hat cf *u<& nor al ^oruls, 
fcat -.ot neariy so broad nor ao io-s ly eo »true ted, In^.de this 
tunic iu the uciaary ee^-cerJ rosier, ore dc-B@ thu u th f~ ale, 
but without or t ; nisaticn i. t : tt.fcuie®, a d re : „ . i .. truit f s 
yovrif .er rule. The rote i located near the* y..e tory a in tr.o 
fe sue, 

. ucr. an extensive a valepxent of i-.coondary c ? -cords . * not 
: ; . , 
uuyr i finitely that ‘secondary sx-eor s never occur in r ts¬ 
ar tins, I his Ci ii,pr-ue:t. a , real contract t ij.e Rt-raul fs.-nle 
in w Lob the* eucc -dory t uien situ : ea ' a lacbi 4 , u •> to ' :.J c:-. 

\ 1.: , f-i i . . ' ~ I:.. \, ■ 

f. ti 1 ■: ■' t 

5ll«rian duct in t * ;.ctio adjacent to t..e n-. in. reduce: to 

{tj ?OrO VOflti. 91)i]C' ti. •'vlft ' ■.; dUCt -O- &51.' t t' • Xv-.r^.-T 

is c rtoinly -ere highly organized than in t & finale (Figures 1 

and 2). ■ |( , 

illier (';>!), hie discussion of th • posible origins of 

foUid u ' Ir the ;: -cord c : -t 








. 






* 











, 


























> 





























































■ 











I section th© possibility that they nay be dorivod fr* a cords 

effect or els© frjiz the -medullary follicles of tr.e ovary* fha 
present case tu.ow.: if At th« f ormer aappoaitio nay h ©ell- 
founded, and presents tit* i•teresting except!on to cases hitherto 
.3escribe • The effect of the rodifjia» > at.ent -stes fr -n? a lire 
after the aec ndary « >*-cords #*re ferae in the fsmls* Then the 
t Ina* to whdeh t > ir. free-r -rtin feolengoi, ware cciXectsd# if was 
recorded that the fusion of the chorion’ ©&,, very narrow» ar.i the 
vascular connection between ther ret-, sli bt* Ifcs bull twin was 
f*7 cr* in length* Th.f p na. cer4itio~? take-, with the s all 

•*#ea the c^orio * wo lc suggest late chorionic fusion* 

I 9*0 <ra. frec-rartin . f, the gonad is- eo.-entiaily live 

fi. i by Chapin (*1?) for her . e®. frec-wertin. There 

/ 

is a single layer of epittfellal c ills c v .-?«i ur4er w* ich 

i? * t in tunica albuginea* This sec?-Jary •• unios i separated fror 
the prisary ore in two rs^ic », Py groups of cell-: the exact nature 
of e* cb the trick-ass of the eootioas «u.de it i possible to deter- 
sine* the rete is located in the centre of the raouila* ffee 
idillerian duct is vestigial. In the ridge on the rese.'-apbrca ad¬ 
jacent V tfceggonad# «hile the welffian duct is better dowelo..-e;i 
than • • • c a ml fenale of the acs» size. 

In .-n 13 .7: c^. free-martin, twin to a 14*0 co* iaal©» und 
bavin.- its head a;.d shoulders a > large as those of its twin brother, 
but hind quarters rush s aller and cv-nparablc to U>o e of a 7 or c c • 
assfcryc» the £Or:ad is tsuefc reduced in else, uni ha at'achsente and 
ducts like those o'- a sale* motions show tw ulstinot regions, 
one next the -eBOntsr j like the rate re. -c« describee by Qhspin for 


















• 

* 



. 




















. 










« 







•'V 



























































!• . 


her It .3 on. iYee-cartia, and another si liar to the cex-oord region, 
tut little? .;iffere? Hated into rde. It i net cXasr whether 

this is u secondary sex-cord region or & part of the odisll* of the 
ori^ineii ovary: but, in any oa.- it ia definitely separated fre-* 
the ret© region by what rwn-bUsc a definite tunica albuginea, it is 
no da "loitoly separated trtv the g^minal opitheliun a: in th© 
re^t c f the , o ad* but it is nevertheless separated fre it by a 
t*\*iie similar tc that of a very young nl.- abc*t 5.1 c: ♦ Tne epithe- 
1 ar. covering t« s ^cmid i. flattened and appears t be *rly one layer 
thick, .the rote has a tec-re. definitely orgaoiiei region near its cent e 
with X •s.nis danse iuiui a© around it-. 

The L o .i has a prisitive ft*idhiy-io-iifco connocticr. at its front 
end, lifc^ that in the sale, and ■*:: ap endix to-’.tis or ©sails hydatid 
of or?;a(. i as a revmuit of the ontlus tubaa abdoiizjale . j firbri&e 
of the ostiu, w ich ha oc-ste connection forearu to the re to of the 
gorai in th - £ nattiery anterior to the gonad itself. 

ibe gonad of the !?• c . free-aartlft studied* ftp roxiratea that 
of the 3.1 os’. Dale in si so and .* coral ap. earance, but wit le-.':s well- 
defined tunica aliupinea* h< *ro arc . o see ndary <s*x-corde. The 
priaary tunica albuginea is thinner than in the 3.8 cs. sals, [he 
primary oex-corda are not as well organised as in the 4*5 ea. sale. 
There is a paler-stained sone between the sex-cords and the rate* -ini 
it is mar this sons that denser growth appears to bo, and in it th&t 
the vaac L-; found* not in the i vuscu< The rote 

is larger than in the sale and ia located near the sesentsriol region 
or hiius* as in the tern ale* The sessile hydatid of orgagni is 
present in aadition to a coiled Tolffian duct foznaing as epididyris 
round the dorso-l&tsral siis of the gonad. 

In the 80*0 crs. freo-isartin studded, the gonad has developed 
like that of a younger sale. It ia sn&ller th n that of the 17. c' • 


















































• * / 





































li> 


free-nartin* but the uif^arcntiation of tbs tunica albuginea* :$ex- 
cords, and rate, if ssore definite. The tunic , albuginea and aex- 
corci region arc rsoro decidedly sal© in type and renew,bio those of 
« >♦! C". sals. *ho rule reaches a sore contra! position than in 

duct 

.,rou? of asaonepicric tubules. The hydatid is not noticeable as in 
the preceding Ouse and has loaf, its oolusnar epithelial coat. ;h» 
blood vessels are still in the region between tbs reie and the sex- 
oords* as in the feralc, not outside the sex-corda as in the rale 
*‘rc'2 7*5 cn. upwards. a coiled ppididynic is present ao in staaller 
sales but out of relation tc gonad age* It therefore a pears that 
in soas cases the cortex of the ovary, as represented by the secondary 
aex-cords* develops in frae-r irtina before the sale bereone begins 
to be effective, but that* in vuefc cases* the hortiono causes degene¬ 
ration cf the cor 33 fro® their periphery inward» so that they are 
not found in the later stages aa a rule, tut secondary sex-cords 
do not appear at all in other cases as shown by til lie ('? ) for 
a ?•?: c»* fr-^e-rarlin. In all oases so far recorded the gorsina1 
epitbelico is cut off fros the sex-cords fey a secondary tunica 
albuf inea like that of a rale ->f a younger cta, e, if net by the 
primary one as in tne sale* This indicates a considerable saouist 
of variation in secondary aoor-coria in the fr e-rartib. In four 
out of the five studied there is no definite evidenoe that these 
hove ever formed* ut in the fifth case (?. osu froe-tmrtin 1, 
fi.i. r.'.) the cores wore highly isveleped and undergoing secondary 
degeneration* his is correlated with reduced six© cf rVse-mrtin 
gonads '3 compared with aorsal r.aloc or feral ea in early stages* 
later jceetiries followed by increase in size far beyond that of 
the cereal ovary* and ac sorption of testis characters with seller 
size than the norsaal t-- ti a. 


















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i 











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"'he TolffjUn iucie arvi Their Terivsttires 


V 


ih? origin of the duels in both oalo *nu ferule• can b taken 
■ * f s . ,. r c • maie /bey 

haw alr^.;.y reached th* circus urc-«*• itai.in In 1 o . ? c* asbryo 

etuoied> at w. ich sta^e ire uretorc n^o easily aeon a dorsal di¬ 
vert acu^a fr-otp the mo If iar. duo* 3 # soar tfco . insi*. o described ic 
the text books for iha PL as other raw ale • :he rectus also c. tors 
the sinus a little posterior to ti e j? ini of sniry of the oiffian 

0 . i ri rined 1 * , . ..r;; : u to 

for a* aP'-roxissloly 1. ct. 

ho s xu&I dirfe'-encoa 1 th .' olffiun x sf proper are evident 
u<- to - *1 C;. , ..t 1*5 cv* the ureters or ntill Joined to the 

volffi r. yets, and at £.:• c: • they c o off these roots Just ahero 
they o en into the Jnui, at rhiefc tire the ureters pa-.n laterally, 
then forwar: fr . thoir point of entry# ana dersally# *nd finally 
Join the central d??ct in the kidrey; at ?* r.., in the female 

they have s para^i fr. ti. > oiffian ducts, an; enter tra sinus 
farther forward# or. .* r. i£ * in M ? icrsal si tsac wall* "heir 
course he the kidney *ill ct be trace - -rtner r’.t 1-. sir point 
of entry into the inis mill be noted a s xuul ; ffererxe* In 
tu. or:-* a la (M, • .;) tfa. core of c un laterally shore 

the ducts open to It. It tn ale th -art of the sinus cc-V aon 
tho '-ani £ of 'i.s ulffiu iuctc ,n..: teo a o ’ 4 he urot "3 
lengtbe-s r - i 1;/, arati-*; the two eta o o>-enir..s farther 
an., farth r* T ita place tmefc a ro lowly in the aulo, so 
that i tee female at 9*0 ce. (fit. 8), th- die*a ce is eariy 
double tr-iit it is In the la. ) e~. da ( . 17}, a difference 

w. ch persists t the '1.0 c=. rtu o. *hicb is as ^ur as this in- 
?dsli 4 fttioa is curriad (i^u es II and 1.}. 




























t 




•> 




* 






■ !*- ’ 






* 


* 






















* 

. 

* 

. 





n 


In 9*0 o'. free- lartin So, 1 (f it * 20), thi distance is as 
Ido. ad, or io^igor than in th© fenale. • In ;So. i>. of the a mo alee 
(F-U,.* ■•! ), it it ?i. r or tba: in the fo .In, a- . n ‘mat c- 1 *© tbo 
^llsriao du ta are carrie s farther hue*: th in - o. 1 or in the 
female before unitin *dtfc the airua, «t 1?*Q c... (Fig. < ), th© 
relative Mata ce.? are f© ale, froa-rar- in, .*lr 6 ©, 2• ; 

and at 50*0 c •• (Mg. ?4; tho &r©;- fetsal*, 4.~; free- »riin, *;.4; 

: - , . ■. . hi *ri • i• 0- t i firiiil HI . c : • is " «c-r i 
in orig in, «h ile *he rest of the 5 -:as i sr,* d©rra . 

Wolf fian duct ft 'r ■ er - r. i: aie. 

At o*l o . (Fir. d), tne olffia -acts s; aw evidence of 
retro-cresoi by a li ht narrowing at th ievoi of the posterior 
nd of the utorus where th Siberian euebs are ce r « .r.i«e. Jo 
••einai venicie prlaordla ar - 4 0 bo aee « 

At ?• css. (FI . ?), the cl fi .n due « of tha fe-ate have 

' 

die inii>hot: in cror.-s-aocti r ‘ t roughest, hut a ir.^lo all at lateral 
outpciiohis.i n each side is u© n in the region bvnida the vagina* 
about Mere. long* tne be roI©< ue o !> the ^eisin&l veh icle. The 

dUC-:: ' . ‘ t, fi 

t e • ?»OR»r-hro:' # where they receive th collect in iuoul or- of that 
orpa end era i distiiif-ii rhabl* fros ire^, to tla 1 sinue. Tubules 
of the r.-son pbros *nt*r the duct a throw,; bout tho whole length of 
that organ. 

A t cs« (Pi.,. s ), the feaalo ej.ows further reduction of the 
7'olffian ducts, but the »rs still continuous, thov.gh c: slier than 
the filler!an duct© throughout. £e irml venicie ar present »» 
two s^ll separate lnieral diverticula fro- ©acr: olffior duct 
about 100 aiora apart* The left duct is the larger in my spebiroo. 
Fror this sloe spv <r < iis ducts ; rso vehicles r the tuft xii* sees 











































































. 





















































to ire a «dv.'= net of tfeccs of th© r: ht ai re i>: deroicpatent, Tb© 
.volffiar, suets t*re relatively «<• .‘Teh reduced t that the? ur in- 
corpora tea into the -.ecen ory later© of t no w-ilis cf the ''Slleriaa 
duotc, lateral •no &•. aswhat ventral to those • 

In the 16*0 c • (Fi, * 9) formal© f the olffia atsc t i prcaant 
anterior t- t* © gonad in t: rs. i<v of th crardal ► rsu. > of to by its, 
but : ..*3 lost -ill conre-C’ :.on to the gonad. It is. indiBlisgisichabl© 
free th© tubules in ap. .arose It has ©cnpleto’y dir ap: ear®d in 
tho region of t ^onaa and a- *ar pcrtericr as th* v-entrul loop of 
th® uicru , where it is r ooe t a solid cord a- -j than ir-ter- 

s tent and v-..ry faint in section r cr a sfc?rt dint© cc, It is sr- 
tiroiy absent in the ration of tho body of the utmis, and ir the 
cer-ix uteri ration, it i. represented by t ro© ©"mil vesicles, 
separate fro each other and dorao-lat ra: tc the c rvix. These 
are the hossologues of the s sisal vesicle, Posterior t the a© the 
duct la nt for a c ratio Hutu .a . Fros about the ant rior 

quarter of the vagina, the iuct is continuous to its 0 >enir;g into the 
eissue. In ao./-e laces it ba? a laoen* but it %-■■ solid for the greater 
part to the place whore the vagina bat-laa to narrow, Fr ~ there on 
it has a !u as increasing is siae, 

At 16, 6 c~, (i., 1 ) al~o, th- ^ If flan duct;:- ar . ist«m ttsnt. 
They are antirely absent in tbs region of the cervix and bc.iy of 
tho utenr , in o rs© plao«, while in that regie the C f ioal vesicle 
hcnologues ap ear as outpouchi .*.•• fro tk* ducts, where they are 
present, the ducts are c stinuoue fro- the a-t rior ©txS of th© 
n©son©-nr or bae tc the body of *n. rtermu h v right duct is ore 
degenerate than the loft, & lun©;.: if proas at only in scattered places 
in front of the brea* in the ducts, and oppo It© tne o toricr ha 1 
of the vagina. 

/it 23#0 ere. (? 1 . 11) in th > feral©, tut olffian duets are 
. iv? ?ntir i, i tr.e r. 1 and '-arts an* 'ric- tc it., and ever; 

t t 4 cranial gmuo of aonepbric iubulee i alrost ns .■ ; nearly 















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ir, :J - tin. i-: all ar act.. >•© au.-t first app -ar-’ n the ventral 
wall of *he uterine horn’ til 4 : « p e© wh re 4 hey b •- v »ftrilly 
and hack iowar tne bojy of 4 ha tiV tir, r a © 11? cord which 
raaohon back to the lovei of the union of the roms, & cr 11 pioco 
of it rsrait :* o • on the side of tbs eer^ix ut^ri ami has an out* 
poochiug of it* a 4 ? a se^ir I v» iclr ho-do oa* iiote ei 

farth ?r buck* o * th v r?tro»l^toral . -11 f *? '• v if inii # a-'-tbor 
1409 t v. . duct an- another a >tslnal vtslola boa * o, ue appear, and 
about th© aiidle c tr.o ina, a third occurs* These thro© aro not 
connected at this r.*& e nor are they cermoctoef to the other duct 
rernanta* Posterior to a rhort ~uac behind ti e lari sertiered 
rern nt, the duct becc-e** c-'etimious* but Ivor off ** l»a t two 
-'.ore little ide notches. Th© duct is solid baefe tc the posterior 
fifth o r the vagina* but ha:; n a lar. irr. 1 nor: behind that, and 
c ; -cns into tne v* tra! • id** - •" V. wa. inn '• ar th: ;tibulc. 

There ere present in the female four or five vest! ial hc*o- 
lopuec of strain*! vesiclw, nous of at ich oc Id be s nr . rs rose 
c auction* ;he ducts, where pres..-rt* are found l- the v&rtral and 
lateral v allo of the va, inu arc uterus in th older tta os* 

■- ; . olffiirn . : uctn in tbo cal s• 

ihe V- Iffian ducts a-o the sa::e in aalea and resales up to 4*i 
css*, but at cl c~. they a-e lar or ir, iu on in aaiec. Prir;oriia of 
sesinal vehicles, c .« c each sid„•, appear dt till a 6ta c .c, an lateral 
diverticula fr--« the duct. (hie. 15) in tbi region the '-tots are 
1 arg -r tb-a- in the ferae I o. (M-• 6). 

,* k t 7*5 c . (’’i . 1C), the v ©t© are till deveio -in*, wails in 
females tney ore not (ri * 7). In th* mic the jcalml veeicl s am? 
propron; in BttS £i*ee out 1 tv, t • n *8** >^1 a no or 4 ri -ri . , fcu 

ur.fcracc ed an yet, a 4 on* or each .lie. The ToIffian duets open 




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separately the & intis • u% in the 

«,t lu # 0 c . C iim 1 ' )* ti ? : tia ' olffian ducts with cc>- 

t rucua luotn of a w;ai variabl .<w-*te at -Jiff rent 1 t©1*. 

The 3^* inal vesicles .-o r* t branch, but, tb®ir h sti ..ate ruch 
larger than tLai of tr. o ducts &t; ja e. nt to the , ar.d *h©\ jevilop a 
thic ' sail l&t rul »a i. btiy dorr. 1 to that of the duct** the 
ducts open to the si u ■•■li '■* ly or-* b l to the va. tr-al ©r->rlr., * 

:?... c itloi of the pata of the t stia to the fftlffUa iwt of 
sao* side throned: the crar.Ul £roup of - .sow hric tobnlcs id *;© n 
in the s Sdi y Is, t- icfc i' reco^nisable a such, 

t-.t 1 ".C* cn. !&) the -‘olffi&n iuot-- in 4 b. .1© bsv® lunirsa 

throopfeo,-* ; the rate of the tt»tia c?-rr sets ie tabula* o- tr ^t oivan 
t. 0 : . - , * . 

duct: ©tr. ‘i t? itol Cf-rd , centalnia t th duct, ha. Ttiicod a back¬ 
ward loo; in the region d.e.ra the ruber?'aculur attache* to it. Trs 
the sale tho trlffian duct? take coot rt in the loo*, ehil in the 
^ trial a tre J*llsrian ;ucts taki at>«i part. In the ado at this tu. ■■* 
tbs ^elffi&r ductc ~c<. upy oet f the cord since th- IfHlerian duct 
c rivativ^s arc alnost f %erte« Tr rales the mri al veniel-s are lar, e 
a n branched, but hav a :>,1? duct lead la.; to to o ffi.r, r.ct on 
each *iido, ir.to which esc! c o s# actually, well co relatively, 
nearer t:© o enin&8 of the e ducts into *hs glens the * -» earlier 
sta es. the female at. ♦ a etu © ha- at least t e or thr -e sts. 11 •«.*!- 
.uordia of ;r;c 1 variclso ©:> e ich side ai i such arti.er forward, in the 
rs; i c whera the due to are i terrupted* 

At tic., c . {Fi . ..9), ths ch&n, ea in tee Wolffian uctc re¬ 
corded above for the 7.0 c -. aala are '-re accentuate-;; • The baok- 
«,-:rd loop c the . © itai cor ? is -*rked art.4 the duct Jartieipate 

in it. r «ioal vesicles are I-r; ;r s\.i ore krone od, a;id ti-oir single 
ducts extend back f.-irthor a~d opo . into the oif iar :vc*% a in the 

























































































!~«0 cr • e bryo. Prostatic vesicle nirro'. • •© th o-lari or ome of 

in© SoIffion iticits &; i uo"Imi vesicle ducto. 

I only one case in all th . al s studied, in co- irasi to the 
feaaler f van ihsr:- any si.;n of acre th& cm. prinordion of < sin&l 

veaiele© © aacn id©# and ihs o.-ly v ry inute aide pocket off 

the Wolffian duct 1 t behin the coin © e. It did r>ot show any 

sigr of ievcio,in into a duel or va icle. 

■n | duetts in the f r .-e-rart n* 

Tr. the •. c . freo-psortirs -c. 1 (>if* 20 )# the lead - edified 
one# the v.oIffine ducts are ©e.-V.nuou. tfcrcdf boat# &» in ho ©ale. 

In the cnitat cord they bear the sn e relation t the 'A llarian 
ducts a?, in the ale, b-:n larger titan the .ftl'leria• due*.a !• the 
a terior part, but svuullcr- in th fo tarior part# aeu in if > ulema 
a d va. ina r&; icon, rise sorInti vesicle# ar. -n }o u .:• unuroRCbec 
a© in the rale# but of only vbr- t ; m -third tba length, and enaller 
in diaseter than in tr.e ^**l« # They barely reach the ease six* of 

lursen as tit a~all part of t'-.e Wolffian da ii adjacent to the ■ • Thi 

free- artin ia r= ah s&ogor t. the ferale type in »i*e rdatio of 
the Wolffian au©t* tlii* t the ©ale, a© fo-.nd in the 1,0 ca* one 

studied, or uvea in th# ?,5 cr • ue* ;h® aee nai von a dec ar© 3 rail 

tut in plto,: of two niwite diverticula on oao olffian duct# as in 
the fmm'-*, it ha only one an in the mle. ha connection of the 
duets witr. the tvboiae of t be &onau *hroujE.h th retc and era ial 
tgpov of '•' Tgoeeidiric i?iiul>.r r i. retains^; . 5 i s&alea* ho duci- j are 
modified feral© ducts, a • t.»e c •• jiiioa seen here ©a- bo easily ac¬ 
counted for in conformity with the interpretation of th • go* ad ir. 

. J(, 

the influence of the eale honsone, &:.v h-n already reach*© a fairly 
def: nite finale differentiation fee fern the he- one co id ecuee re- 

i*c 1 i on i n th '•■ 1 e *i l roc * i n, 


Tbit: ”-ii© r c. * on is -err: evident in 












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tho anterior parts the ducts, than in the posterior* 

9*0 oa» freo-aartin 3o* 8 (Pig* 81), which has a ore rale type 
of genad, has its solffiai; duets else clecer to the rmle typo* They 
are an 1-rge as the t&llerian ducts, beside which they lie in the 
region of the uterus and vagina, excepting the posterior quarter of 
the wa>;iaa which is larger than they* the seminal vesioles are 
larger and longer than in Ho. 1, and even show signs of a branching 
in their anterior ends. They are larger in lusen than the ?olffian 
iucts In flat vicinity, but hawo c-iy one duct ••etc,-. :he olffian 
ducts are ooaplete and lit.'ed with well-defined epithelium throughout 
even though the lumen is very minute i thoir ant rlor ends* In thtt 
part of the genital cord -interior to the attachment of the ;;afeer- 
naeuius ligament the Mullerian ducts are absent, except for a small 
piece nearly opposite the centre of tho gonad, an a piece near the 
ostium tubas abdominals, which ha not completely de er.eroted* The 
ffitelffian ducts extend t the forward end of the gonad, where they 
are connected to the rate of tho onad through the rsHiwuntg of the 
cranial group of Wolffian tubules* 

In the 11*75 cm* free-aart'n (big* 88), the Wolffian duct on 
each aide extend forward only to the point where the guberaaeular 
ligament Jcims the genital oord. fron this point forward, the cord 

r 

is present but has no lumen, nor sign of a duct in it* Posterior to 
this point* there is no lumen in the aenae of a definite epithelial 
lining for a c nsiderabl^ distance, but it is marked in section by 
an area of degenerating tissue. There are five asperate little 
evaginations of the Wolffian duot in the seminal v?siole region on 
each side, a female condition, and of those the so t ant.-rior one 
alone attains any considerable length* Of the t® ducts, the left 
is the smaller for some distance anteriorly from their opening to tho 
sinus, and each has a luron as far forward aa the sesiMtl vehicles 









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27 


extend. fee ur.tror seminal vesicle has a larg r las o than the ducts 
thesselres, and shows som oi^na of- preparing to branch at its anterior 
end. In this ease the Wolffian ducts show little ef-ect of the rale 
horacne, and were presumably so far advanced in festal® differentiation 
that they were already d generated in cose regions, before the ss&le 
hormone started V- act* But tha development of th* antricr casinal 
vesicle is a sale characteristic. 

In the 17*0 cm, free-nartin (Fig* 83), the Wolffian duct I®, 
continuous froa the front of tha mesonephros to the region of the 
uterus where it is interrupted In four places. The left is better 
developed then the right, but both have the interruptions. They are 
again interrupted in the region of the anterior seminal vesicle, which, 
though one of four on each side, develops in the true sale fashion, 
and :?v«n branches at its anterior end into tubules with a coat distinct 
free that of the genital cord, but like it- The three posterior di¬ 
verticula on each aide, prisordia of ucoensory seninal vesicles as 
in feealos, are sacra narked tha*: in the 16*6 os- female (Fig. 10)* 

Of all these nominal vesicles, the anterior one on each side is 
siailar to those of a younger scale, and the lent three are sorely 
imperfectly stimulate - growth centres held over from tne female 
period and not inhibited completely as in the sale* The lucen of 
the posterior part of the Wolffian duct extends forward to where the 
anterior seminal vehicle cornea off on ouch sid • In this free-mariifl 
we hmve a^ain a c ndition int«r»»diate between that of a female and 
that of a younger Bale. The supernumerary serlnal vesicles are 
cheroot ristic of females, whereas the ant rior seminal vesicle 
eondition and the continuous aatrio** end of the duct ars distinctly 
sale* 

The 20*0 cm* fr e-aartin has the nost extreme modification in 
the male direction of the five studied (Fig. 24)* The ?olffian ducts 





















































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are oontiaoouo throughout their entire length as in a stale* though 
the lusen is not ac large nor ®c well carried forward* The 
^fillorian iuota are repro anted only by a short, but patent utertia 
sasculinua in the region where the ge ital eor s separate* but 
attached sore to the right cord than tc the left. Lillie figures 
it on both sides in soegc of his epecimtna* 

At the front end the ducts extend forward pat the place where 
the.r last oennecti n to the Wolffian body io cosplete and branch 
into minute ducts* that load to re corbel parts of the olffian body. 

Tho eesinal vesicle© are anoh acre highly oovelopod than in tb<» 17*0 
cs* free-isartin. Thera are no leas than eleven of those on the left 
aide and about as nany on the right. Of theca the ant rior is the 
longest and thickest* and hranohen only near its tip. The s cond 
ia next la length and a little r*ore branched* The third is a little 
charter but sore branched* They get progressively shorter posteriorly 
and less branohed again till the last two or three are n rely short 
diverticula projecting vcntro-lateraiiy and then extending forward 
along ha 'olffix*n duct. The lumen of the duet narrows Jest 
anterior to the opening of ths anterior a ni.mil vehicle. Tho femle 
of this else ha only five snail octpouohing# and they occurred on 
an interrupted Wolffia s duct. 

Here again ie an evidence of a female oendition later codified 
in the smle direction by the -ale hormone eh cfc stimilaied the 
Wolffian duct. The mle at this siee has but a single saainal 
Vehicle on each aide with many branches* all opening into a coraaon 
auct* that passes back and opens into the olffian duct near the sinus* 
In the fer-ale the seisins! vesicle hoesolcguoc are farther fross the 
sinus. In the free-r.artin tMs distance is intermediate. In the 
fcsale and free-martin the Wolffian duct o each old© appears to 
open into the floor of the vagina Jurt b fore it open© into the cimia. 


































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tihiX* in the aal* it appear# to epen directly into the r> nos fr * 
the aedial sides cf t«o grooves lateral to a s dian ridge cn the 
dorsal since wall* the froe-wartin resembles the feaals more *ban 
the sale in this rempwot. 

To tnrs tip the ftolfflan duot situati n it say bo £ id that: - the 
first sexuxl difference to appear is ia the relation o ureters to hft® 
ducts* t?p to 8.5 e*. t>;e ureters ccao off th® Swiftian lusts in both 
9 jX85* At 8.$ aa« the ureters in the fsraJLe cc>-« off the sinus an¬ 
terior is tho ducts, while in the male they cose off with the duets* 

The distance be tw e en the o?«ni ge of the ureters and duc^s cm the sinus 
wall t eis loryor in the ferule than a the aale and is twice as lcn& 
as In the &l e at about 10. 0 cm. this s *ual difference is maintained 
through the 80.0 ss« stage* foe fr#e-aart3o ia generally intermediate 
in cmditlor, but ***re naerly approximates the fmal«» 

The female has a aarrowin; of the ducts throughout, ap earing 
at about 5*1 os. and progressing* till, at 15* > 6®. the ducts &r*? 
interrupted anterior to the vagina re^^n, and entirely absent in ti e 
gonad rep,ion in s we oases , but present and continuous in that regien 
in others. This depencration p r oceed s till the ducts are present only 
in th^ utsru-; wal?, as solid stripe, and in the walla of the vagina, 
as isolated sin t* sag-sente in the anterior half ant ecntinuou: in the 

i 

other half, with loam only in the posterior one-third* 

In the male, tbsse ducts ret.i & connection with tbs tubules of 
the gonad through the cranial gr up ~seonepbrle in ule* and rete 
of the monad, which is early lost in tin female. The ducts continue 
to develop and i have a lawm ie th^ir anterior end*, and occupy 
the whole genital cord in ports where the CftUerian duct is generated* 
In the female the Banner of opening into the sinus differs from 
that In tfco mole, in that the ducts Join th > vagina before opening. into 
the sinus in the later stages, *fcile in the ssale they o en separately 



















































































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50 . 


to the sinus. 

In froe~~ariins the conditio?- of the duets is T&ri*blc. "She 
eenfisciiio of the gonad tubules to the Wolffian riuot is reta csd is 
four of the five cases studied# *r.d, in the e caption, the whole 
of the interior pjiri f the duct was ^{taorttai a . in the f conic. 

Iha ducts tisasilTss eery in cone it! on* but, with the e 'caption of 
the Il.tfc cs. free-sari in, where they appear t; bar® fe.hawed aa a 
female cot for sous ccncidarable time mad degeasratad# then to have 
bean stlaslated; they are me c in typo. represent V oca of a male 
of an earlier stage. 

Ae to the aeaifi&l naloltt;- tboy develop s sevitat later in the 

female, 7*5 cm.. chile in the male they appear first at S.l css. 

In the fanale there are fire or perhaps nor# prirordia that begin 
to develop at various stages, but none reach *m? else ap roeiable 
in gross dissection, ana in later atagaa they appear or; an inter¬ 
rupted duct, end ssmstiae* an isolated parts of it, always farther 
forward than in nalea in later ntages. 

In sales one vesicle develops and grows strongly tc a coa^ider*- 
able long!?, before branching, and its single duet epos* into the 
Wolffian duct progressively n&urer the poster.or end of that duct, 
as tbs erbry© increases in si so. ?» ©as ease there we-* « eiinuto 
second outpouching of t-.e solffUra duct in the region eher* a second 

vssicl* wot-.Id cone in s fatale, but it did not develop* 

In f r ee-ssart l ns there arc an easy me c oven amofe prissordia . 
that get a start, -hey are not so lon^, a- in rale* of oowreapending 
else, but are developed after tor. jaolo peiierr. In ihi* also there 
is varieties in free-asariir*. of the sam* also, here also the 11.75 
es. freo-ssartin (fig. 22)# is like a fcasl* Van tfc« other * and 

appears to have developed lc- ger at*- a female. .In l.a larger otag.es, 
though store mssffQCs than in the male, they develo like then. Ihe 
oldest cue with its eleven separate vesiolss on each side give* a 
alee picture of a gradient of differential growth and inhibition 





















































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ulon.- ti\Q axis o the olf fieri .uct i:; tp assaina1 aside reg on. 

In the froe—auriin the openi t f ti e a -cts nt- tns sinus is 
sor.et- '3 direct, tu. i t. a ils, .me :• distil. : t t; v. va,;ins first, 
as ir. the fer al®, .hi.. t..-ar; r r: i<i 4 iDi. * t< e ctat® of* ether orgars 
in refer® - c® to tr.e decree o infiu :-rco of * he rials horcon®. It is 
3ore lik tho rul.• ??: or® V.e other part, are ,.cro affected. 

Ii.o free- art !u ciffian .net derivatives j»o® pro, roorive modi¬ 
fication o- a feral- ; rly oi ’ fsrantiaiio toward a c^ie later dif¬ 
ferentiation, under t 5 inf. icnce of a ;i ie ho men®. ire a te ree of 
this odi icati ■*. *p- : ears to dmpG’.d on the tir e of inception of' the 
horr.0- o and tr.e time • i la o.- r. * at inception to tfe sts o s*u::ied. 

, c ulloriiuj . acts ..i.- .‘heir derivative . 


T-. tho indifferent sta^ ;- tr.e Siberian jueio fc .n to devalo? 
at stout ." c • an a . li-r.t thick®r;i\ c of *i e co^lon ® ithelium on 
t e v^ntro- adial surface of the Wolffian duct ri(do t one- 

luarter cf the ;ji + a c tack frois the .t-orior an: o ‘.he mesonephros• 

but no inva inati r. i tc fe<. -n. , t !•* cm. the or- ties* tuba® at- 

jo* 1 ; .ir at;.: ir.t r.or and : f th-. tube haVv ee fc u •. t- f.rn. .ho tube 

develops by a. infoldin* of tr.o ccaicv:. vail on trie v tro-lot ral 
surface of tho mo aphres ju t re .trui to the olffiau. iuct. i’roc 
the first i 4 lini- di f f sr.* fr the ccelc e ithallus ar.i from that 
of the o.-tiun, Ih. e iti. ilii. is colu-’.ar, but not n thick a that 
of the ortiur, h.r o tiu li n .tout one-.carter of the sesoaephros 

lar.i tt back fr its ant rior end as : ®r ; schickir (*1 ) finds in his 

1.: cm. e tryo. at 1. on. the loot appear at it pe terior end as 
an infold ir., or thickening of tr. e: * ei.xl trail of tr.e olffion 
duet, an.; >:* in-.h; for ar 1 bo ter® -n thi fall and tho cooloa & itfeeiiua 
as a distinct cord of a diff® ant *.Itheliu , more eolu nar than eifcho 
of the oth re. It ha#: no or ; . anissd lum* 


in i v . po t-r-or ... n ■ cn the 










































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32 


left side* w ioh is in advice of the right in development• Ifee 
cavity of the ostium extends forward pa^t ito opening into the 
coelosic cavity as a pocket with wrinkled walls and the coelom 
lining is al c t ickened for a short di ta~cs in that direction. 

The total length of the duct on the left side ia 83d cdcra, on 

the right » 374 siora. 

At !•? on. the oooIcib epithelium anterior to the o tiun 
thickens raark dly on both aides. Then cn tracing the sections 
caudally, a fimbriated slit froa the cooloraic epithelium leads 
into the interior of the flat duct, at about the eaae level on 
both aides, lined with taller columnar cells and lying ventro¬ 
medial to the Wolffian duct. This ends posteriorly an before in 
a cord on the ventral wall of he Wolffian duct. Left «ide is 

855 micro and the ri*ht is 705 ricra long. 

At 2.0 os. the ostia are attached to the diaphragm and the 
ducta have lamina for some di :tanc*. the left being greater as 
before. 

At 2.4. 2.5. and 2.7 cn. the ducts show increase in length 
in which the left is in ^dvanoe of the right. Ho such difference 
between the sexes in the setter of one or two ridges for the 
Wolffian and dfcllerian ducts, as ic described by Felix (*12) for 
the huimn enfcryo. is found in this carles for cattle. The ostium 
is farther forward on the right side than on the left due to the 
backward migration of the mesonephros as described by Ber*scbickor 
(*12). The ducts do not reach the sinus in these stages. 

In females:- At 2.8 cm. (Fig. 3) neither right nor left duct 
has reached the sinus tho gh the left has got as far bock as the 
opening of the Wolffian duct to it. Lunina are present 1 1 rcughout 
their length. So separate Ufllerian duct ridge is prose t in the 
fe-aale of thin stage. 
























































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33. 


3mJ era. (Fig* 4)in ferales, to ducts reach th© sinus poutorior 
to the oiffian ducts* but h&vs not yet united to f or ■ uterus or 

vagina. Tht epithelium rounu the ostium io raed^ura columnar, 

/ . 

At 3.6 cm. (Fig. 6). the o tiusi epithelium extends from the 

wfclleriar duct to thsjpto of the go::«d* where it reaches the coelor 

/ 

surface, after joining, a gl ntrulua of the Wolffian body, rhe 

iihlicrian ducts of the two sides cosk# together in the median plans, 
but do not fuse to a einclo lumen, and noparato pojieriorly before 
they open to ths s*nuo on the aedia side?? of the dorao-lotoral 
grooves* along which they extend for a distance before their lining 
disappears. 

At 6.X on, (Fig. ti) there is a slight epithelial* but not tubular, 
connection of ths ostlu , aith its tall columnar epithelium, to the 
rote of the gonad and a glomerulus of the eo^ono^hroo. the Uftlleriar 
ducts hare lu&ina thro- nout but sculler than the Wolffians, except 
at the anterior end. Posterior to the point of attachment of the 
round ligcnent the ducts enlarge greatly to the borne of the uterus, 
and are four to six tines as arge as in tha saloof the same oise. 

They arc beg lining to have the transferee portion sore mar ked and the 
left has more backward migration than the right. They unite to a 
single luaen in th© uterus region, then are double again* and again 
single in the vagina region* The Wolffian ducte Join both the rag ins 
and the oinui at about the same level, but the vagina projects back¬ 
ward frosa this po nt of junction as two buys with a tongue of tissue 
between then and join ths sinus separately from each other bat not 
separate free? the Wolffian ducts. 

At 7mb cs. (Fig. 7) the ostia ere joined as mbove to the rete of 
the gonads as well as to the gftlierian ducts. he left cstiun is farther 
posterior to the right than before. The rote connection is wuch smaller 
than before, and the ostium lining ir. rauch teller than in preceding 

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fesiauea. ib© ostium is non as far forward as the tubules of the 
as son© hro~, and the ‘nJllerian duct extend© farther forr&ri than 
the iolffian arts, which In represented by & teklt in ih© tie,7 ©ae¬ 
rating Bs^onephreiu In v* re - the ^llorian inct is 

throe tirea the sis© of the d©fere rating Wolffian duct, arc is 
actively developing. The uterus horns ar relatively larger and the 
corpus lit or 5 end vagina are forced *& single tubes with double 
regi ns between the®. Ar. Judged by the epithelial lining the 
utillerian ducts ‘oin the 3i ss behind and separate fr the Wolffian 
ones* 

At 9.0 cb. (Fi^. $)» tb© ©etiu has turned to face the gonad 
anterior end* and has a partition down its iddlft* and a ciliated 
lining* The details are well described by rierpeohicksr ('!£) and 
will not be gone into hero except as they relate to sex d:ffarencea. 

At 15*0 ©». (Fig. 9), the infUsdibulua is seen as a little side 
tube off the ostiuss, to the rote and tine to the ©md, but it does 
not oenreet to the sex-cord j of the gonad as is the nale. ''he 
whole connect!'-n is the epoophoron, parovaria- or orgar of osanesiller* 
as contra* ted with tho ductal! efferent©© of the rale. The coil of 
the uterine horns is basoning »cre aooectu .tod by reason of their 
own growth, net fro m a pull by the round llgBoent* which is attached 
half way down one side 'f the ©oil not at its tip. -lie annular fold© 
at the ©or tx appear and the vagina, with its sooth lining in 
contrast to tho wrinkled lining of the uterus, enlarges greatly. In 
its mils ar© found tho rsanants of the soslnal vesicle hor^ologuas 
or artaer*s ducts, together with the Wolffian ducts thoBselve*. ihe 
posterior end of tho vagina i filled wit h epithelium, and the Bolffiau 
ducts join it, and a groove from tho vestibule sects thc-v radially, 
instead of two *.rooves ae in earlier stBges* The vagina extends 
posteriorly as a dorsal corridor of the nl^us. 








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he 16*5 os, (Fig, 1G) nta&e shows only an enlarged v&gina* 

I 

and at £0,0 cv. (Kig* 11)* the whole genital cotd is filled by tha 
.Fallopian tubes and eliHai horns and ihj ©oil of the uterus la 
complete* apparently aa in tb adult* The rote still joins the 
posterior end of the ostium. the vagina ha:; enlarged tremendously 
in lusssn. but net in wall thickness. The vestiges of the Besinal 
vesicle honologuas are in tb thick walls of the cervix uteri and 
anterior part of the vagina, Ahe dual origin of the vagina is still 
evident at its posterior end where it io still divided into two back¬ 
ward projecting bays in ite dora&l part »b i i ~ the ventral part opens 
to the vestibule aa a single tube aft r receiving th© olffian ducts 
as before described. 

In naiesi- The *&lleria& ducts are porter in naloc, smaller 
than 2.6 cm* (Fig. 1?) than* in females of like eixe and do not yet 
reach the sissus. *hero see a t be a sexual d foresee in the rate 
of growth or formation of then# ducts even in stages that appear 
sexually indifferent externally and in which there are no inter¬ 
stitial calls in either sox. (Lillie, *17, *23; Basco®, ’S3) 

The duots roach the sinus by the 3*G oa. (Fig* 13) stage and 
are in all r© ;pects like the feisale ones except for want of luster* 
in the anterior and extreme posterior ends* -nd the fact t : at they 
appear to open to the Wolffian ducts where they are open t > the 
sinus. 

The ducts arc solid in r.my places by the 6,1 on, (Fig, 15) 
stage and show signs of de ©n ration. The connection of th rate 
testis to the ostium i© larger than in the female. The remnant 
of the duct is in a secondary ridge on the genital oord, The ducts 
fuse to a single ut rua ra»ouliana but smaller than the female 
uterus. They are double in the cervix region and single in the 
vagina region back nearly to the sinus* where they become double* 
and join the corresponding *oIff ion ducts* then both join the sinus 









. 

<1 u j 














» 

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: - 

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round the tongue of tissue, as in iho f«oa3,« t but the MSUerim ducta 
extend farther fc&ek than the Tolffieeo, 

By ?.» os. { ig* 16) the ducts are discontinuous, and the eatiuia 
has-loot its thick lining of columnar eoltheliuis, but the ret# testis 
ocimects tc it. The hydatid of ttcfgipi is present ao a «atU knob 
on the aiae of the ostites* The lips of the groove that fuze in the 
female to gi e the Anlleri&n duct, have not fused in this specimen 
and a slit takes the place of the duct without the characteristic 
lining. Snail re casts of the degenerated duct or® scattered along 
the gonad region in strong contrast to the feral® condition where it 
is actively d veloping. Utercn. ua culinun is still fcrnm. ^ervix 
in solid, bat a luaeci reap cars once and is gene before the vagina 
proper opens cut with its character!, tic lining. This divides to 
two b fore opening to the s bus as in earlier stages, but here pros¬ 
tates era visible in section near it* 

By the 10.0 on* stage (Fig. 17), the e tier is practically gene 
as such ana the ducts are gone back tc the gubermcular uttsfliami 
to the genital cords, where the right is found such better -reserved 
than the left, one has a luram first where the cords unite. The 
Uterus masculine is about tha sis® of one of the olffian ducts. 
the dftlierian derivative# arc ebse i altogether from the uterus 
sasoullnus to the place where the lolffian ousts nearly reach the 
sinus, where the Vagina waaouliim appears, almost filled with epi¬ 
thelium, and dvidea to two, and iherse open to the sinus, a?? in earlier 

stag s. 

At 17.0 m. (Fig. ia| tte dSllerian duct derivatives occur only 
as the isinut© r.ydatid of hergagni, Utsrua *eccelin»s '4th short, but 
wide lumen, and appereatly a derivative of the left duct, and Vagina 
aasculina, with largo lt&cen near the ainao* awl cucrtmnded by pro- 
static# tubules, Its Unit g is nut that of the normal vagina nor 








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is there any opening; or connect! r. to sither Wo Iff ion ducts or ainxw* 
At 20.0 cr, (Fig. If), no vestiges of the vBllerian ducts ware 
found anterior to tho region of union of the goitai nerds where an 
unpaired solid body extends bac' tc the surinal vesicle regie, dorsal 
to the Wolffian ducts. In tha region of tfaa opening cf the .olffian 
duets to tbr si min the vagina a&aoullns appears as the rasruuct of the 
rikht 4fcllerian duct* rroundod by prortaiic tubules* a d connecting 
with the dors 1 wall of the r$,ht Tolf ian duct, posterior to its 
openir.r to the aims. It la solidly filled with s'libelI uk. 

In the frea-isariiBj- 

for brevity tha following terse will be used for -he different 
parts o' tha ^llcrian duct derivatives; 

Ostitrs fimbriae * for the ftmaai-liV* anterior opening t the cooler, 
when acre Ilk*:* the female condition* and hydatid of ^or^ag :i * when 
like the nnlc. 

Tubal po rtion * for that part that corresponds to the Oviductr c t 

: allocis** tubes. 

_ . _ ~ i - 1 

Corn u* uteri for the parts that corres; end t; tbo enlarged transverse 
cornua of the female. 

Ucrpua uteri, for the part cor -ca coding to ito body o f i 1 k ( i adult 

uterus. 

Uterua nasoullnnE * where a derivative is partly -efcrable to both of 
the above, but not definitely 1c either* 

Cervix uteri, for the re on of narrowing just behind the corpus. 
Vatina, for the part cerr sps ding to the adult vagina in the female. 

. ■ . .M*i. m i 

aeina eK*oculina , for a condition that approaches that ir» the ®ala 
where only the pc- 'torior ends of the tubes are present. 

In the 9.0 oau free-raartin do. 1 (Fi*. 20), the least modified 
of all those sto led, the duct ia intersittout in the tubal portion, 
and continuous behind the resonephroc, and roserblo^; & stage inter- 






























































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V 





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se. 


between ^ale? of 5.3 aci 7.5 crs., bein^ X^as continuous than 
tbs forrer and ^cra so than the latter, The esticn# consists of a 
*bmU knob or rid* e eof jr«4 with ooluamr epithelium and * slit Into 
tbe neeonephrcs anterior tc the #onad. Ihia retains sore of the 
columnar typ© of epithelium tiian the snlo, but does not clor.e in to 

a a i&llorian A fio .o face the foaati an la tbft foaole. 

However there a e sifens farther posteriorly that it did ©lore in or 
row backward, and later degenerated, in the shape of a degeneration 
spot at the baae of a very snail secot dory ridge on the Wolffian duct 
ridge. The latllorion duct has no trace of nierior opening tc the 
coelom found in the fordo, but rear the cstericr end of *ho gonad 
it hs* a Ivm n quite as larqe as the ^clfftan duct there. The cornua 
uteri have enlarged ti I larger than the Wolffian toot there, but 
wfcc-rc the goaital cords unite the tftllerion are xssch smaller than 
in the foaale, though fo radar, a single corpus uteri, acre Icrsal in 
position than in thr fenale. bo cervix is oingle, *he*-e&s in the 
ferale it is double, and it ic single throughout the vagina region, 
the vagii.a joi c the odnus round the eaaa tongue of tissue described 
for fe*s*ile specimens, together with the Tfeifflaa ducts, but the 
va.- iaa lining do=?c not bocor.c included in the sinus ti 1 posterior 
to the yoXffia^ ducts; this free-aartin so id appear to have had its 
S&llerian duct derivatives devolved to a fairly udvanoed itago before 
degeneration aas induced. 

bi 9.0 en. fr>i ■— rtln 80 , 8 ( ig* 21). *ith e aach reduoed 
gonad *ti ecu arecl to do. 1, has alneat an little dative ievsiepaent 
as th$ 10.0 cc. sale (Fi?. 1?). The Pierian duct ie object in 
front of tbs aiddle of the gonad, at which point it appears, enlarges, 
gets a luseii for deal era, I sap pears not to 

reap or till rear the junction of the round ligament or fe ;ui •rncculu’i 
to the genital cord. I: this it closely re cables the condition of 












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39. 


ih© hllorian duct n t.hu aai© , i 7•© ox Ij.O os. froa it* point 
of reappearance it is iK>r© o ntinuous than to the 7*4 c&. la^l©, & 
o ntra t t© it» condition ent-T- or to that point# It ic praccnt far* 
ther forward on the right aid* than cn the left, and bos a iuceu on 
that aide before th© iaft appears, ihe cornua unite to for' a single 
corpus uteri, ehi oh c r. ,tricta to a carvix, about one*third the si«.e 
in lucin of the corpus, «nd than, as the bagirm* dorsal tc the olffian 
ducts, it enlarge.;. greatly uni flatten© out laterally. Posteriorly 
it fills with epithaliu and Joina the a.mis, uu in prsdediag cases, 
but bacorea continuous with the ainua as two grooves in the dorsal 
ainut: wall. 

In the 11.7& cm. free-martin (fig. ZZ) both Yolffian and ^ll©rian 
ducts are discontinuous, Jae rat© connects with tfe* coelom a* in th* 
fasale, and a hydatid appears at the end of it as in *h- sale* do 
ilSHorian duct i* found anterior to the ooreua uteri *.ed then only 
in part of in© left sid©, eery fa nt ona intermittent- the geaital 
oorda era .each reduces ana tho Slletian ducts ars both absent, just 
in fr nt of the union of f hf5o* Ocrdm* Asphericai c. tcri makes 

its appear a.-o’; for 200 rsiora, with a wide iuacn, a remnant of the 
left aSllerisn duct- a vagina s.eculina a© cars, both in tne riddle 
va&ih^ region, and in tb> posterior vagina region, whore it car* a 
wisU lucen, but fills with epithelium Just anterior to »ho a it Joins 
the sinU 4 , after raesiving the o>*tti pa of th© holf ion duota round 
t*.e ocuitossaury tongue of tissue. In tills region this embryo is de¬ 
cidedly farsaia; in the anterior rogiom- neitaar 2 ial© norof© --ale. It 
appears M tiio differentinti: in lb Wklm di--©t- • non not 
starter till after the fnlffin ducts hod d»g*n rated very decidedly, 
but had started aeon enough, und acted ion& enough to cause vary 
threat 4*g*o*tf«tEUB of tho MbUtfiM due* l-r they reached a 
ccrBiderabl© development* 

In th*- 17.0 ers- free-oartin studied (fig. 23)* tho ^llerian 




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40. 


duct is di continuum at the anterior and. It appears for about 
160-200 ''lorn a* % solid ping in the ©eseriery bosiia the col lei 
Wolffian duet that forsss the epldidjels in this spuci? n, and 
also a • hydatid of Morgagni rsore definite than in the sale of 
this oise. the cornua utori of the left aide appears at ih* attach¬ 
ment of the g^ersaoalQRf but does not extend forward into the 
lateral part of the tranr.'reroe portion of the cord, but on y back¬ 
ward, to .join the larger right horn to for© the corpus otsri with 
its large loaea. The cervix ia solid or interrupted* ami the vegin& 
is patent in ih anterior part, then f lied with ^pitboliura* but 
patent again for a short distance in the posterior region, thou 
again filled with e pith slims. It is eery evidently de# eneraiing 
throughout. It receives the Wolffian ducts before opening to the 
sinus as in females* by a dorsal .roore in ih - median dorsal sinus 
wall. Thereare no pgo ciot — bar# as in the sale. 

In the ?0.Q cr. fr s-e-martin (fig. £4), the ostium ha* failed to 
©lose in anti i* not found, the condition tbor -t iu like tha^ in a 
sale between % 5 atw id oca. and no hydatid is found. 40 Ifelleriar 
duct derivative is found in the tubal or cornua regions, but its 
former presence is indicated by a s all ridge on the aeaonephrla 
ridge* k reranaat of the corpus} uteri persists as a patent biird 
tube or reside in the wall oftha right genital cord just anterior 
to the union of the cords. The vagina la like that of a 15.0 oa. 
■sale, coctio ed, b t rot ft ured for this paper* but a little ~or© 
degenerated in the eiddle region, and relatively eligbily larger ic 
the posterior region. £ber? is no sirn of cervix uteri nor anterior 
vagina re ion. The posterior va*in* region ie such larger than in 
the tsale, an receives the Sclffiac ducts before it ©pen* to the 
sinuii as in the fewele. The luwen is wide anteriorly, but filled 
with epith< liucj p© tcricrly. ’to prostates are yean arouir it. 

* 

Discussion* 

WM>«H rntmm m 1 i mmurntm*' .- 

In the case of the ihillerian duct derivatives, the conditions 




41. 


Quad in ihs freo-aarttn a -tt easily o<ar«!ut#l and explained, if the 
*5llerian ducts carried on a© fanalo til they had a ooi .tart is 
development, thtfu b«o* H, « sradattlly inhibited &ni recorded> under ine 
inf-uerce of the naie hormone. At ih c -a the horson* ot-r.u- 

lated the active development of fee olffio. duct ; -riva* iv , ar 
ahowo in the increased length end sice of the Wolffian iuc*e mod 

-»*i v ' ' cl-": , -.. ; •' ‘h~ ^IfUOM 

on the eirnts fcetwe n ureter opening* and those o r ¥olffian ducts, a 
fesele rtiirilt-r, coupled with -everal feminal vesicle prirerdis. as 
in the fecaio, activated, is produce enlarg d reside* comparable to 
these of the sale. Id eo i» asotraed here, the sale and f©- ala eon- 
dittos are sodificati ns of a ecsrsen sot of Wolffian a i ?&llerian 
duct and gonad priaordia, in two different directions* by gyf.otio 
influeno a in the female, and by yy^otic inf', cnees increased by 
horaone action* in the male, as ft. own by th-.r above details? then 
the free- nrtin, Judged by the ea-* criteria, 1* « leva loped 

ncr&ally by the early octir, cyretic influeneea, up to a eta*:© which 
varies with the i dividual ease a:: c ntrolled by the s -called ac¬ 
cident© :h©rIonic fbtloa ar-d tiiwllf ! s^anm * edi¬ 

fied in the oale direction by the sale horaone, ascreied by her 
twin brother and passed to her in the atoral blood tramftasion, oatiag 
a retracing of the path of fe-male ievslor-mnt, followed by develop* 
sent in the :aale direction. 

The ^aocus Vaginalis, Rou nd Li gage nt sr.d Quberr^iOulicr . 

for purposes of d scrlptior, the tern Scoeua vaginalis will he 
used to designate the cavity or see that develop© free the vaginal 
process of the peritoneum in ih> inguinal rc gicn, in both males and 
fesaias, cad inguinal cnn^l its part in the regier, that Icier close© 
in adult sals. The tor . dsnal of • Jfuefc will not be ua?d. ■uberua ou lus 
will signify the ligausni of attachment of the genital cord to the 
bottom arxi wall of the aaocus vaginalis if that li^asent thickens and 






























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42 . 


lac (IftUHMt la the ftfedo* I . c. fcy#aa6 dot; ' penetrate IqpilllS 
region anovo iV-rpart's li^-asosi or it& ^ni valent. la fyuo^aeartisa 
the two conditions c^y Apply on tht different aides. 

Up to 2: *6 on, the saccuc a:>; iigjajsort are practically identical 
in the two aax-38, Xa tfco case of the foualo the coccus vaginalis 
usually do.a not enlarge relatively after thiv utage* sr.j though ihero 
appears to ho s a variation in this re. poct, it n ver approach a ta 
sale condition, in which this sac groan out through the Icgniaol canal 
tc sufficient length finally tc reach the bottom of the carsium. 

Neither doe a the v&ll of this esc thicken in the feral e tc the sane 
extent ss in the ale. In the ramie the ligament lengthens and thickens 
is ifc:; wbdesisn, then shortest, while in tha ma e* it does so in the 
saccus vaginal *-©# an.. tfl a greater extent particularly in stapea above 
20 • C Cui • 

At 3.G cs* (figures 4 and 13), tin femle has a etuillovfor euccus 
vaginalis than the said ana a slightly shorter lipanrnt. £y the 5,1 
ma, stage (li urer. 6 and 15) the formic has lean tfcar half the depth 
o? saesue found in the male, tho ;$a the well is not noticeably thinner, 
the iigneeat is losgsr in the abdesea,. and leads wore laterally than 
in the cale. trie is show; ;n gross dissection fe tier ibua in sections 
or recons true tie-: figures , or, only I erso-ventral dlaplooeeenta ea- be 
aiovn. In both .;axes, tae ii&asasai grows do^n inside tho coccus, at- 
tacaed along one side by a aeaeatery (really retre-peritoc X at first)* 
and to the hot tea. 

At 7.5 o». (fi * 7) In fecales, the lig&sent has elongated, 

broadened. and thickened in th> abdoaen. The saccu« cavity Is no 

do por then th wnll 4 in its solid botto extent a ittlo farther 

into tho tlesu? «f the Iflfgin . I regi n. The direct!. • growth of 
«t» MM* »*U i • -. po 1 -rc-li.tw«l •■**■**. Ir. On 

guterrjicului)! is osrro»**r in V‘ s abdo-.^n tt*n in tho 

but th» qrocUw cavity i. abent two *al or.o-half tir.ee ao deop. 
































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43 


and *he wall & little thicker. Its direction after entering the body 
wall i nr o v ntr d. 

At 9.0 ok. (Fig. ft). tte rorud ligament ir thiokoni). h in the 
abdomen of the fersale rore than in fh» 10.0 oas. rale (Fig. 17). Its 
dirooticn is SMBdad -oro than wenirad in the body wall, vise aaccuo 
depth has not incrsasoJ materially, a nci the ligament thickens inside 
it at the a awe rate na in the abdowen. In the 10.0 oa. pal© the. liga¬ 
ment is tninner than in the ferule, »nd tighter stretched and straighter. 
The aaoous is ineospi te in the propartation , but is vary such longer 
and deeper thi - in the female »v*n thou h inoonpl te. she i uberneotil un 
extendi: down its sail, but, the direction la acre vsnirad than in the 
female. 

In the lb.O cm. feral© (Fig. 9), & mark'd ohange i? eviaent. 

The round 1 iganeirt in the abdomen is noon broader and thicker than ia 
earlier stages* am the cavity of the aaccus is relatively end probably 
actually ancrtor than in t • on* stage, but its vail ia tb id 

its direotion ia still lateral, eno caudal, not ventral. The genital 
cord has taken a decided ventral loop in the regior Jv t anterior to 
the attachment of the round ligament. Che widening of the peiv.e, 
rapid growth of fha uterine horns, anterior to the attachment of the 
ligament, the retreat of the f orad, end the failure of the seccus 
va^lxutlls to p not.rate tho inguinal ccnal, bav^ contributed to keep 
the growth of the ligament, ell in the abdomen. Tn tho lo.O as* oa.e 
(not figured) the -ubernaculua extends alnoct directly froe tire nuen 
reduced genital cord tc tho body wall as a thinner flap than in the 
fenale. It is inolua^i rolled up in the inguinal canal, and instead 
of becoming solid, as in the fenale, *he suocus ©xtorids ventrally and 
caudally aa a long narrow pocket with the ubermte ’loa folded inside 
it am attached to itc dorsal wall by a sessnt^ry. the ventral margin 
of the gubernacnlu* is thickened, apparently paralleling the thicksniag 






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44. 


of the round liga^e-nt in the ?s~ale. ?ho vaccuc cavity «&c over ten 
tiiaco an doop -b that of *h* female- of th» »*«« jIxo. 

In the 1-5.6 on. fop.itlo (fig. 19), the round li. a»er$i is thick¬ 
ened in the afcdoosn. The sncoos ic about on*5-thirl a dvtv. at* in the 
I'., or. sale ( v i.> IS). but i-> i* th r- 5 r no thieketiir -vr-gp ndi ru. 
to the ^uhernaanltni, nor i» the :^cosi wall thi V^aed tm -och. The 
direction of growth is nor© lat'srnd srd c oadad than in th© le, r.ot- 
wi theta ndlng the appearance in the i Hr at ration wMch is oo-oi»bat 
distort ad by dir«©otion. the « 2 ao«e*.l depth of he *oocno cavity in 
t'is female ahers that there ic ccnoiderafcle variation in the normal 
fo -ale, but the range in the fret-nuriin far evens taps those bounds. 

In 17.0 on, Ral e (Fig. 13) the sacc^s has a large isni of 
growth in valla ©ad bettor,, ’he ^bernwauXusc ir ev#o sore thickened 
ic the aacoue then the r© nd iigiuaeci of th*s fossalo ie 5 o iho body 
ISfltyi and it in ri . : ‘ thicker | n the body cavity tear ir. 

earlier male eta o, eau«i:s.. & loop c : ' thy genital cord, ’.he hotter 
of tr:o f.accur. cp road- "C the asrotu; it t.rally. 

"'he 20.0 ce. female (Pig. 1") ban her round ligasent rsry tnioh 
in the abderea, forning & thick fold between the doreu and ventral 
linbs of the loop of the uterine horn's on each ride, with e a hit-lire 
cavity in it, th ? * origin ©» rhlch 't was l&po£slbls to 'etentice. It 
ic. cot open to th ceelor. as 1st tb> aaccue, but aeeoe to develop by 
delaainaiion ic situ. Th? cavity o? the eaecns vaginalis has die- 
appeared, and the- round ligascn 4 poise tfco broad llggaeot laterally, 
forcing ft thir. Hap c* m I stery a£os* fh body wall V. »-re Helm 
but with no raat thick aalo# in the body wall sh ro it -stars. It 
Just spreads out Info the surrounding; tiayycs ant! disappears. 

In ths male at 20.0 cm. (fig. 10), the ubomaculuc; srterda 
slightly van* rally froa t ha s all ■ a .ital «erd to the a ecu, c tnin* 
in the body wall. It is thicker and rclati ftiy efcorVar than in 
earlier stages. In the saccun it ic thio'-e ing anc leas folded. The 

0 

saccus extends doeoer ventrally toward the scrctu-, alcost reaching 

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45 


its top. 

above details it appear* that the gubernaoulun thickens 
tb..^ tn roii.r. -t ttdl CtC| , ~ l s r . .■ :*H Am tfeC re a ecus'* 

is great, while usua ;y ns appreciate!* osoone appears in the formal©, 
and for this reason the great thickening in the feaclc occur a all in 
the body cavity, while that in the nalo is th=^ s? siccus* The increase 
in length in tbi fe^^alo, and the attach -rant to the uterine herns nearsr 
their union to fora the corpus uteri, causes a loo in lh© uterine horns, 
in addition to the spelling and loop of tbs round ligament between then. 
Shis ligament aeons to shorter and tfei out in later stages in the 
fenal©, in the sane manner ac the ubornac lun shorten© in drawing the 
testis intc the inguinal canal ami do*m to the scrotal s c. In any 
case, it doo^ not retain threu h later stages its relative jsixe a *5 
compared to the cornua ut~ri. 

In hree-caartina: 

In 9. 0 co. free-ssart in 80 . 1 (Fig. £0), the llgurtent is not thick- 
nod in the abdooen as in th female. i relatively a little longer 
than in the ssal • but bands slightly ventrad as in the fenale* The 
genital cord also t~«os ib© ventral bend characteristic of tb« feoale, 
apparently dee te tb£ lengthening of ths cord, no 1 to a pull by the 
ligament. The coccus vaginalis is deeper than in the fenal© of the 
came ©i*e, but it w^llfi no iheker, and its direction o-uudad as in 
tho fodo. While tteis specimen is int^rsediat© in condition between 
th<t saxes in the round liganeut, it is still pred -inantly feral©. 

The 9*0 on* free-nartin ISo. 2 (Fig. El) has the gubornacuius 
slightly thicker in th^ chionea thar c the normal calc* ©act longer, 
and lens direct fron the cord to the coccus o e ing* The direction of 
tb© naecuc vontrad, somewhat a® ir< the sal , ani the bottoa of the 
sac cue ixs denser, thicker, »nd more actively growing than in tho 
feaale. The ventral loop of the genital coris of ihs? femle is wanting 
here. This frtc martin in so © decidedly sale in conditio than So.l 































mm <# met Af • ••■■■■■ : 
















































































































































and ha* a guberrmculua instead of a round liganent. 

tbs 11.T5 C”. fr o-aartln (flu. £P) has it* round li| ament 
thickened ani curved i the abdrren, ouewhat a* in the 9,3 cm. 
feral*, but the curve in dorsad no 4 vontrad as i- the feale. .The 
coccus vagina is la at le al ten tines 23 long an in the 15.0 esi. 
fenale (fig. 9), and three tl^es an long as in the 9.0 era. fenalj. 

Tho ligament extends to the bottos* of this see as in th * uale, but 
it not yot thick nod in tho sc, ncr is the a»c wall nu h thickened. 
Tho direction of growth is vontrad an in the wale. This freo-rartin 
has ~sal© and fewale characters 3 : ©what evenly b lanced. It suggests 
a fairly advanced Resale c&nd tion reversed by the ction of the rale 
horron*, takin., effect late, by roaevr? of late chorionic vascular ' 
anastomosis. 

In tho 1T.0 os. froc-isartin (fig. <?3) tne /uboroaoulua ia re¬ 
latively lon t . in tho abdon~n, cut only slightly thickened there, and 
has a double curve instead of being tight stretched, the moouc ic 
such longer than appears in the figure rcuchin# almost to the udeer 
prir ord ium, and the gubcrrjaculua is attC.n thicken *j in the uccos aa 
in the isale. The wails of the aaoctiG are thick ana the direct on of 
tha active growth ia ventral. This la doo'.dedly rai in type of do- 
volopaect, but ha evidences of feeale early stance. 

The 20.0 c*. free-sortin (fig. 24) has a very marked thloheoing 
and broadening of the gubernacumn in the abdoaen, but lass tj*on in 
the fe~ale. ana farther lateral, toward tho body wall. It fora* a 
decided bulb in th~ ody cavity intsral to tee a-bilioal ortu iea, 
but exte da sodialiy tc its attachment tc genital cord, a a 

thin sheet. Tho eaecus invades the body w&li ventraily anu caudally, 
ab ut ftac-tfeird as far as in the sale of this sise, anile in the fa- 
aale non* at ail is evident, hero th© gubornaculuc thick n* in the 
sac cur and this thickening is w ntinuoas with tnat in the uhdooon. 
















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the saccuc batten is growing nearer the udder regi n# but no scrotur 

appears. 

That this structure io a female om being sodified in the sale 
-traction is shown by the tendency to thicken in lb body cavity, but 
also inside a long aaeons vaginalis* 

An interesting o& e in this connection is that of a £9.) cm. 
fr?e-martin, in which the lsft oid? has a toetis already drawn Into 
the saceuo, and a guberaaculu?. thickened only in the sue, while the 
right side, on the contrary, has a round li.anent also t all thickened 
in the abdonen pushing the gonad farther tack fren the suocuc opening* 
This ceeas to be male on one side and fosaalo on the other, if one does 
not examine too oio ;ely. but on closer examination, tiu > gonads are 
seen to bo alike « cept in ,osition. ffee anomaly ic due to the failure 
of tha right aac penetrate the inguinal canal, and develop, while 
the left did so* and eo one s d developed in the female sooner ard 
the other in the sale. thif. can to understood when it is cocu.idersd 
how far the left i do, u-n a rule, is in advanoe of the* right in de¬ 
velopment, particularly of the male characters, the stimulation of 
the vftdrnl sac caused it to reach the inguinal canal on that aide 
before it clo-^d, or cau ht the left pricerdiua at its critical stage 
and caused the right to grow too slowly to pass the closing canal, 
or missed its critical eta e. In rany frao-mrtins tb sacs fail to 
penetrate on both aides as shown in Lillie*e (*1T) pap r and in many 
they penetrate on both. There would appear to be a ti~e when the 
penetration by the eaocuc i; inhibited* if the aectts hu.i not already 
passed a certain stage or point in the inguinal canal, the loft sescc 
to pass this stage earlier than the right and to bo in a condition to 
penetrate core often than the ri,iit • 






































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r\ *<i * # ■ ' - £ ~ ' 

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fat s of fJhta Ite^cse^hroa 

In the feaale: 

Thd nnsOBephroo reach or Ita maxim* sice In enbryc-a bn* risen fi.i 
*ai ?*5 et*. Tor a*. 7,5 on. it is nborter than at 5.1 c~. ?©rgschic 'sir 
(* t) says the ^axisrua i err ©ached et approximately 6.5 cr, n'>ek-raffip 
length. “e-orptien appears to be greatest is an .:nt©ri-neatcr!~ior !i- 
reotlon* but eerso coers to go an at the posterior end. The ret^^at of 
the eortad any ad! to the appearance of eo terior degeneration* 

Bergechicker (*’?) ahoiss thor* art; fluetuatic e in both cexog* and Ibis 
study did not nsu>.-; comparison** of -snlrmln of the ease ego and sex. 

by 9.0 or. (?*i§. T) in the ferale. the nesoneohroe »hov« considerable 
dearea e i siso. and tMc takos plac© principally by the degeneration of 
the cranial group f tubules xmd loneruli. 

At 15.0 c». (Fig. 9) th» female show® a reai reduction in the 
nuncer ef lcsseroll lairing a aas of tubules and a fee eloneruli. 

The rote ovarii eti.il passes out into the neeonephrcs and leads to a 
gloasrulun, Put no evident tubular ccnncctio'. either to the glouerultt* 
or to tiiO Wolffian duct i, presses as woo noted under th 1 * tftllerian 
duct development, tfc rota leads on to the option tubae abdcri al®, 
but its significance was ~o% clear from thia study. 

At 16.5 on. (rig. 10) in tb female, the crania I group of tubules 
are not retained in any c ip -city and the whole sesonephroa has undergone 
further degeneration* 

At gu.O on, (Fig. 11) the rote ovarii extends anteriorly from the 
gonad end roaches the ooelo^ie surace. forming a fe* tubules opening 
into the body cavity, and covered with columnar epithOiius en their 
coeIonic edges. Ibis Joint th o«tiu« anteriorly, but bar. nc. connection 
vita esomphrio lillwlfl or duct, and Mil out posteriorly. So glome¬ 
ruli and only a few email tubule resnant« are present in the ssee ©no phros 
itself, forming an epoophoron* a the tuoules c nnoctod to th© rote are 



























































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interpreted us r mat? of tbs- ant rior |fo«p, ard a paroophoron, 
consisting of a few tubul«n? at tfcs posterior end of the jtsCnad. 

In ?»#-. 

•he condition pereUels that it; ftttalos tip to S.* on. (lig, 14 ) 
where the esaonsphros &.jpoar*> larger in diasatar, but not longer, than 
in th f.anaie, /it b.i «S* (Fig* 21) it is longer and the cranial 
group of tubules has not degenerate*' as -men as in the festalo while 
tiia caudal grestip ha*. A© it* *nt«ricr -art ic larger ia diaseter, 
tl)Ki ia the fttsala* B1 diffo---o-.ee etc wore mx at ?.h «b v 
(IS > ) tai« K& , C * . (fig* IF)# a aal# has it leas do e r.? * e.; 

iricr *nd than a 9*0 an* fitale (Fig* 8), bet Jsat as such 
in tbs caudal group, £i 17,0 Con, (Fig, lb) tfc# art rirf or cranial 
group cf tubal $ s rsuai-s 4 i foir-a part .? the apididy-is corona© ting 
the rcte.it' tia directly to tho iolffian duct, bat ih ? caudal group 
has deteriorated ani glcaerdi arc found only an rudiments in the 
paredidyai or orpin o* .iraldo* ^ad in •* ductus: ahwos. Dcctuli 
efferents# are forcing a connection of be Wolffian duct with tfco 
rote tubule# of ihs tostia, but the glomeruli of ther,e cranial *»so- 
n/rphric tubules arc gone. At F0,0 -.tsu (Fi#, 29) this connection of 
the duotuli efferent#* from *olffiaa duct to ret# teatia io plnin 
and easily traced, £ucfa connection v *a? lost in the ftsftlo, wher^ 
the Foil led out to lb * d I group of 

aeacnaphri© tubttlos bs: forced a paradidymis, an a dueto* aberrere 
8**,c n : a off 1 self in th* HiWlir y with three email er»Jarf#W atI 
or. it- tubule/; i * dK.oc rate loner^Ii. Th# w-ol: -•■ -toner*)!*? ie 
slower t a . : r.cratt in the nal# than in th 1 ffcmrift* especially In 
th# later atftgos. 

In tbs free- art n: ** 

In the 9.9 on. ftee-eartia So, 1 (Fig, £0) tfeo aL-eonephro* i# 
larger than in th# fe ale of the auso ai&e. Ih> caudal group of 
tubules are degenerating a littl# ©or# slowly th* in the fecal*. 










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but the cranial grup retain th ir cennedtion to ti e rote and Wolffian 
duct as in sales* llomjrull arc present in both gr-ups of tubules au 
in the aoraaia of both soiob, Thi.. free-nartin resembles the salo in 
longer Btesonephroo and conaction of the onnial tubuios to the ret# 
of the gonad* It roe. nbleo the female in baring the re to lio'ced to hho 
ostiue* but less defi itely so than in the feralc. Ifc© rote is not y«t 
organized into dvfirite tubules* 

The 9*0 oss. fres-rartin £o* 2 (Fig* 21) has ei cre reduced cranial 
tubules but a total length greater than Id feralec* The o&ud I group 
of tub los 1c not so shrunken a In ;,o*l, lb?? connection of rets 
and cosloe it enall rr t;. |jb the feoalo, but ilia ret;? is c msocted 
t the clffiun duct by the cranial group of tubules and glomeruli 
are preaent. 

The 11.75 on. free-®artir. (Fig* £g) haa a sari?oa c n triction 
of the scaonophroa between the cranial and caudal groups of tutu los. 

In one place there in but a single tubule in crone-taction between 
tber. Ib© cranial tub Ira per at without any connection to a 
Volff i ri duct, which ia wanting in t i - re ion of t‘is free- martin* 

The ductuli effarentee a a ; rm t *nd h rate is ccnnectt i t the 
hydatid of Korgagni or osti -a rornant, and only to th« posterior fee 
tubules of the cranial group* Though iso olffian duct co id be dia- 
t ogulched from tna tub les, it probably ©si ts coiled upanong thee* 
Posterior to the gonad* tubules are acre numerous and a fe» degenera¬ 
ting glomeruli ary to be found, but n~ « ! oiffiaa duct• ;he Wolffian 
body as a ml cle is almost as s all as that Of a 15.0 ©a, fermlo, 
but degenerate© in the solo runner, aft r losing its ducts ao such 
ac a 16*0 c~* feral© (fig* 9)* 

the C3. (Fig. fid) froo-sartin has a such shortened meso¬ 

nephros in which the cranial group of tubules sees® to be involved 
in the retreat of th© f pmad» for it« epididymis and th^ car al group 
appear dorsal to the era ial ones# be ret© har> lost its connection 
to the hydatid, but retains it to tho crauialgroup of tub los and 
thua to the Wolffian duct, Sloaoruli are ab.-ont* ther ia sore 







































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mesonephroi retained than ia eit&er male or female* but tfe:> relative 
ptiilti<:*n vtsa c'nna ti -?ne of parts retained are with slight modi¬ 

fication. 

In the SO.C c*. fr*e- artln (Kit;* *’41 th ret© rotai * it* con- 
noction to the anterior tubule reruns in the ori -idycio. but boo lo©t 
it with th hydatid -hich is also absent* * dtsetoa oborrat!* 1# r*pre- 
3 enteu by a aigle isolated tubule, ext-rndirg for - ara fr •:.-•■ th'? nsepne- 
pbros in the body wall, ©operate from tfc* ether tubal*** Jlomeruli are 
•beent, %ilc th© cranial group of tubules is not yet $uite ar» -ueh 
stimulated *p in th© male* wticro they for* the dootuli efferent** of 
the epididymis* there is nor© aesorjepiiros as a whole then in the male 
or female, 

the degeneration of the aea< sepfcres in the fn*e~steriin Ac clower 
than in either sea normal* but in general* it follows the ale uethod 
of disappearance* in that the cranial group of tubule# form ducts 11 
effer ante© of th© opididycia. 

The grogenltal Tinuc and It Derivative© » 

An before described* th* separation of th- urogenital witaua 
frorj the roc tun is complete by th f,4 or, sta a. In the female the 
peri nets- remains short* and the urogeritf'l sinus o ©ia to the exterior 
through the urogenital plate at thtc early stage* while in the wale 
the elms remains clos >d. ar*© a ridge cr raphe ferae vortrally over 
iha uro ianital einurt, which in prolon ed to the tip of the fiallue by 
a narrow vertical plate of ceils derived frer the urogenital plate, 
the per.ia r.igrates root rally a Ion? the belly wall to it© definitive 
position close to the umbilicus, This greatly olo-gates th* urethra 
In the mils* particularly in the fmm -ereranaooa and pare cavernosa* 
and directs It antoro-vertrally, instead of posteriorly as i;s the fe~ 


sale. 





In fbealoe at £•# os. (Fi&. Z) tho urogenital plate, * lch cx- 

tenda fro* the perl nets* to the tip of the phallus, is perforated by 

<& 

the posterior o© tinuution of the urogenital aimie lesson in the fors 

of a narrow elH to the exterior, attending saso ilata no® ventrally 

fron the perineuc, but leiving; tho ventral two-thirdf: of the plate filled 

with the epiibeliun of its walls. This plate extends along the dorsal 

surface of 4 h- phallus: to its tip. the prissordia of the fculbc-vestibular 

glands are already present, as o all latere out*»pouchin&e of the lining 

epiteliim of the two dorsal-grooves in the side walls of the s-rus. 

Bhil* only about 100 nicra long, they turn forward as distinct glased®, 

but are net sc far dorsad as in tho wale of thic siso# end are slightly 

ssore apparent. Frcr these * lands forward,.the clous, flattened ©u* 

laterally, bends all htiy war trad, and a ridge develops in its dorsal 

wall, extending farther into th luzsoa as one proceeds forward, till 

tho sinua itself hocc^es & narrow crescent in section outline where 

the Wolffian duet;, open into it. this they do by penetrating tho 

inror mj.ls of tisi" crescent from tho interior c the rid,? e. the 

* 

l&llerian du ts dnot reach the sinus at this stage, but they do at 
3. ' 03 . (Pig. 4), Just posterior to tho Wolffia ducts, the distance 
oo the urethra proper frewa ureters to olffian ducts is longer in the 
3.0 ci5. rta s. 

At 3.6 css* (Fig. 5). ths ctnua s a little longer, the bulbo- 
vestihular glands larger, d the sex ducts open into a sinus, oval 
in orosa-secticn. The urogenital ; late atil extend? tc the tip of 
the ph alloc, the po ter lor vulva opening has an: urged t» all ii~ 
mmionc , asd is : open for alnost tha entire depth of the plate, 
the dietaoo ofl Ih© mi sthm ureters to the Wolffian ducts, 

is longer, th© porfneu*. i~ a little wider, and the phallus, a little 
Tsoro recurved, than i.i younger stages. *h® - ttlc regions of growth 
behi d the vagina are now incorporatec into it. 

At 5.1 o • the urogenital piste extends forward into the tip of 















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t he phallus an a pouch, fi .lei «ith aplihallua, arid roofod ovor by the 
recurved tip of the pialka» which flutter**? out at tha 15.5 e». stags • 
Tho poriaous widens a little by tbs 5*1 oo. stag*, (Fi... 6). but 
thereafter it just ko^pa pace- sith too growth of tho surrounding parts 
«nd > oto relatively narrower fro the H.5 c?. fttage on. The vulva 
enlarges so sishai, but, while at 5.1 oc.. it entert i:i as ant ro-dorsal 
dirootioa, its floor gradually flatten? out usd by the 15." or., stage 
(Fig. 9). it leads acarly horizontal y into the vestibule, vtile at 
20*0 <ua. it is horizontal (Fig. 11). Tfco vulva ke p pace with the 
growth o- the surrounding structure®. 

The bulbc-vestibular {leads gro* oat laterally fro* the cider, 
of the 3 nut where the vulva leads into the vestibule, and they begin 
to branca at the ?.£ as*. ota. «? ans there aft r boeose ncro branch c-1 and 
?5«aii larger, till at 15 .f cr, they penetrate beyond the saccular coat 
of the vestibule, r-ncio «<J is a ouacular coat of their own air covered 
by the cr rordiusr o <: the constrictor rnlvue auscle. they o ntirue to 
lacre so relatively to the 2%i e~. at&ge (ki... 11). Their duet? opaa 
to th" veatib 1« far 1 h r vantrally c» the sides cf the v ret i bn Is than 
do thair ho ologuaa is the siala. 

the Y-«tifeul* bsoewts relatively nbortor end larger in lunsci, 
and becomes alias t double by reason of tvs ri;g.*a, era in each lateral 
ivall, that almost s. rpurat a longitudinally wrinkled derail corridor, 
which is lai&r stages l?ads atraig. t for*nr int the vt; ins as* ir? 

15.u os. fossa 1* (?ir. 9) frors a ventral corridor, loading, ferrar to 

the urethra proper. 

/.t 5.1 os. the vagina and WolffIan duct© open intc the sine®, 
one behind too other, from ths -ails of a ridge in the dors**! sinu« 

vail, rotmi a tongu: c r tissue a* described under ?4*ll*rian duets, 

\\ 

asd the vagina baa two baoWard projecting touches st It* posterior 

before it. 

end. By 5.0 c? 3 . (n^J. n) the Wolffian ducts .join th- vagina o •■ms 





54 , 


to the voetiend at the 15,0 or, sta:,o tl*• vagina opens straight 
anteriorly fr » the dorsal corridor of «h© vestibule as in later 
stages, 

.,o«c&lied hoaclcguoa of the prostates, the p;*r*~ur«tfer*l ducts 
d. «Ot Mil their uppear&ooa tosffcr. ea. stag* (Fig. 11), *fctn 

ti^oy appear ac two t r throe snail aide duo to off the dorsolateral grooves 
of the rest-hole# a «;hcri dietarnse posterior to the vagina orifice, 
they do not develop in the poet.'ro-a»ter ior direction oh rnoterieiic of 
tbs mis and thv-y are such belated in appearance. 

Anterior to the v&gino-vestibular opening, the urethra proper 
•steads forward to th$ ureter openings. As described before, this 
distance length esc rapidly In fee fezsaie to heoosje double that in the 
ealo by the 3,0 ee, stage, (Fig. S) an,: still longer relatively, later. 

At 5. on, it p&ssee forward without narrow.ag noticeably, chan, as 
froe a square cross-caution outline to a crescentic one, and develops 
& ridgo in its dora.il wall, through which the ureters suiter it. Ja 
later stage©, whoa the suburethral divert!cuius develops fr-» S.O ©». 
upwards (Fig. 8), it hfia a sudden decrease in ltmen, Just in front of 
the vagina opening, with the auburethr*! divertic Ins developing ac a 
forward projecting hilad poach, ventral to its narrow posterior end. 

This divert 'oultta i nor eases in ais.«, and finally, by the 15.0 os. 
stage (Fig. 9), it feraa *m anterior continuation of the venire: corri¬ 
dor of the vestibule, with tfe? very narrow typically female urethra, 
opeaing into its d roal vail near its interior end. This diverticulum 
enlarges and lengthens up to the £0.0 ea. stage (Fig. 11). This saasa 
to have no hoscloguo in the sale. 

In ’aaloor- 

The uroge ital plat© doe. not beco&o perforated as in the fesale, 

* * 

sad, at £.8 ca. (Fig. IS), its edges in the m ddl* and dorsal part have 
begun to grow together to fora a raphe ventmlljr froa the pari us. 














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la thj a^rsui surface of the phailne, it* Heroins nasi the akin are 
cio appreciated tfeaa ia tho feaola* the afeol© of It is fillod 
with ep--.tfee.i-. .4? tci Ife., per ia os i - feet sli btty brewer dorao-r*. - 
trolly Ua» -a the feaala« .*u 1 bo-iar *tbral £la d pariaordia are prasoni 
feoaolo^iia • of tha bulbo~v* ti'oular laadn, but a.-v ~e d^rsally placod. 
ina luaaa of tfe sinus anterior ife th^se glands ia nor oeer this in tfeo 
fondle.. Ihs off flan ducts onter at it - anterior end* feet tfe rid.e 
tjQfmz. m the f Jt*ale io wanting, -fee '^lleriac duct* hav* not yet 
yearly reach©I the sinus. Ir. the apeeiaen studied tfeer? va* a -^aU 
blind lio i with Solffiaa duct tpithel. ur, caudal to the rip,* 

KOljffi a duct, in the dorsal ainuo <*sll, but *•* it do*r * ct appear in 
later cta c .. a it ia taken to fee of no **xaal el^nlfieeeee* the ureters 
j-ais the eider- of the olffiaa da-.-tn ad sinus ot their point of junc¬ 
tion, and the sicu* extend* forward as the urethra or neck of the 
allantois, without narrowing. By 3,0 c% (fif. 17,) t the urogoaital 

ainus reaches the surface only os tho dorsal surface cf the phallus, 

» 

and ia filled with eplthalius tercu^nout*. Buibo-urethral , lards are 
rore developed than their hoqologaao in the fecal*. The t:nu», anterior 
to these ducts, ban a larger dorsal ridge than io the feral®, and froa 
it the £& Her lea and olffian ducts Join the si tm% together, but the 
JHloriaa® extend farther feacs as two diverticula. The urethra proper 
is shorter than in ifc j feral e, but hr cider, the 3. c r. dale (Fig. 14 ) , 
has tr.i following ofean-■.•»<*;« the migration of ths p nis forward along 
the wastrel body wall and "hr widening of the perineu her continued, 
the bulbo-urethral |luad apertures have roved ventrally on tfce else® 

jus an bar a pocketjibe enpra-ur- Ural 

dlvcrt-c.v. I? ? c- it fir; c : Lly Ur*rci Uic rectu , :he 1 •-■..! of the 
pro static oretfcra ia 1 than in the fetal©, and the v p llari&n 

ducts join the Wolffian ducts, and the i&tt r join tht sinus, but the 
ridge in which they lie it relatively smaller then is earlisr stages, 

Th© e x differences at this stafetf are;- the r»o u-ved unperforatec 





















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cavernous urethra, the rsi^ratod penis, the dorsal pouch In tbo 
fculbo-urathral re ioa, the score weatrai bulbo-ursthral glsrda, the 
lecher proatatlc urethra, the short nidor urethra proper, tad the 
aider per mews of ths sale, as oc&pared with the fesekl9« 

At 3,1 os, (?i&, 15) the vibration of the peris past the serottnft 
has lengthened the Cavernous urethra, which 1.-: narrow<?r and perforated 
so® disturoo fro® tbs prostatlc port tcw&ri the penis, the corpus cavcr- 
nosoa urethrae is developing along its arters-dcrcal surface, and but- 
rourde^tll but its ventral surface. The bulbo—urethral glands under the 
prinordiu® of tha buifcc-caverncu* roscl* and the supra-urethral divert!- 

oalus at the bend, are longer than in earlier stages. ’he pro t»tic 

» 

urethra ha; narrowed very nceh juct anterior to the duots of the bulbo¬ 
urethral glurda end prostates firnt appear a a snail solid diverticula off 
tha dorcal and lateral grooves of the sinus, called the prostatic croovss. 
ihoy ep-ear first at the posterior end of the proetatic urethra and spread 
anteriorly and circumferential! in lathr stages, The prostatic urethra 
is rel lively longer and narre^or thar in the fbnale, The aHlerisn 
ducts open Into the sinus cep rat©ly posterior to the Wolffian duets, 
a d the urethra proper in shorter tha in the fenalc, 

fcy the 7.8 MS, etsr © (Fig. 15) th urogenital plate rersn&nt has its 
definitive posit-oa t the under surface of the gland penis, and perfora¬ 
tion of the cavernous* urethra in alsost coapl etc. ho si*, acid flexure of 

% 

the pools is beginning to derelo and the urethra traverses it* The aupra- 
urethral diverticula at the bond in smaller relatively. Prostate* aur- 
rcuoi the pr c urethra and extend forward alaost to the vagina open¬ 
ing, and the bulbe-urethrdr> begin to branch. Th^ urethra proper in abort, 
and the pro taiic urn thru is about the ssu&e length in scales end featnlcs at 
this ta go. 

At 10,0 os. (fi IT), the prepuce begins to develop round the glens, 
and the whole penis points sere anteriorly than in earlier stages. The 
dorsal divert io-ltus la larger and the bulbo-urcthrols longer and ffiors 
branched. Froststea ar oor© profus© and extent nore around the elongated 
prostatic urethra, which is narrower tha; is the 





# 


57 * 

.■ 

i 

fersale, Vagina nosoulina an,-' Wolffian duetn Join It separately# end 
the prostatas roach the vagina c-pcnirv. 3© subu r e thrai diverticulum 

develops Its tho mlc, but tho urethra arrows (interior tc the vagina* 

At 17*9 ca. 18), the cirsoid flexure of the penis is wall 

developed and the cavernous urethra Ion,. Ifec fxoatly enlarged bulb©- 

% 

urotb rtv 1 flands open into the dorsal diverticil cr, at the b?ai of th* 
urethra, "h© pro tatic urethra i;: three ti es as long a- in the fersale 
end irrowor, end the pro ;iat«a e rround it and the va ;rs. h© urethra 
proper is not narrowed as in the fe&ale, and is o ly half gf lan*j. "Tie 
ref ins do-# not ope to the -.inns# but ic bilni, both enter crly and 
porter’ orly. 

The perforation of the urethra reaches the avrt&CQ i tho P.0-,0 
cm. ale (fig, 19). All the a*le character© a e nor© advanced except 
the supro-urethra 1 diverticulum a* the band, w icii in small*-r, Pro- 
state© are tailor dors ally, but few, trail, and scattered, voatrally, 

in free-eartloa:- 

,fee 9*0 oa, frae-aarilii *»o, (Fig. 00) bar a set of feus&le uro¬ 
genital itiave derivatives u ich ccrrccponI to t oae of a aomsal feo*l© 
of sossetrhat .younger «ta^», the floor cf the vulva and vestibule is 
flatter and tne v« ttibul ia short, but not ;uite o larg* in diucater# 
a* in a fc ale of Uhl* siae* the perigee i * in the fe; ale. The 
boLbo-rstibci l«r gl* tda are not .*© far aivanced, The Itoiffiac duct*f 
join tne va;.i a, then both join the aiaoa, as in the forale, The 
urethra narrows in front of the suburethral diverticulum Gad ir even 
longer than in the ferule of this ci*e. Ho pro bates aro found. 

There ir nr- r .-sc.fclunce to the sinus derivatives of tho sul» ic any 
particular. 

In 9.0 oc. frse-rartiu So. 2 (Fi£. Si), the urogenital siaus 
derivative;* ar^ also fecale in -v ry respect# bu © u i .h.tly reduced 
size. It ov-»‘‘ .- &u the sasse *1 all pochat of *pi jifillc*. u>u <or tho t.P 

of the ^5 lane olitorldi*, ac veil as th fe ale vtslv* opsnin ♦ The 


cv 


i 




fwtihiit is snort, »?** in oi&seinrj balbo-v©-tl-lm**r glrcds nr® 

f^a *3 in tyo*, but I sa fenrandhod and a ul or, than in * 3 o. 1* itewevsr, 
ifc* vat'ioa mod £oif£i«tt ducts Ojwrn to the v n?tibu!a e-ap&rat#Xjr like a 
ssaio, Jhs ur,tr.r& pro -er i? twice a# lo g a 1© tho 10.0 ve* sails 
(fig* 17 >» ho subtir Viral di rerileulun ir. not davelorrd, fcrt tfe* 
israthr?- rarroee after tha far.ele fssbioa* Forinear *s ?ir?.* J e tr* type* 
tlie »J-rv* d* 4 * In *h£w that, of th two T,C oo. frev-rair* irs 
stvdtad To. 1 is »-re adv-.need ic footle efcaraoteru than To* f, i* res* 
pact t? ureters, oiffi-t© duct ocna^eticm?, .nd v* inn* mil aesodevoa) 
dleritr lv 8, hot neither 1* in sny re pool ©ale In tin derivatives of 
the urogenital sltne, * ich is derived fr « eutsdvrn* 

75 ■ :o has a tlftteH 

fnal« type of ferlceoFK, vulva, -and vestibule* (Tig* £?) *elb©» 
vestibular gland« are sore advanced tr...n in the 5*6 co. fr -t-oarrtire 
or normal fv^aier of that ai.se, and the vestibule ia shorter. ~s 
pro static fccwsclogacr are fe*,fi4* ’a irs arid olffl&n duct* unite, then 
open tat tr*-'; sirut m l.n th f-.v^le* Th re ir» qo- <?■;finite ubrretbral 
iivs lieu lost, but the ur; thra .ounwa in fr-nt of a dip in the vertibul* 
floor os in 6 isasfc ynunr.er fe.'-ate, and .. de/^er?*r»t$ne fei ’o® in fteurti 
dorsal to the eatsri>r end of the ventral vestibular corridor* where 
tfc§ urethra vcela b , if the suburethral divrrtieelm hui developed* 

This loohr, a.- though »©»e d*» enerfcii© cf & feaale o'n'iti r? were £©• 
on• The urethra i- trice a lor* a in a ?*$ Or* f«oalo and four 
ti sc «e long *© in a l 1 *a o * L-&X&* hi fre«-«orti-i ie also fe sis 
In sinus derivatives, hut l*h r~. <1' ioa* 1 us in h s direction in 

the .-eeodernal d rivative , ife? Junction of the tnrihra to the subure¬ 
thral div rticulur or veetibuio* 

The 17*3 *n, ffee-®*rtin (?ig. ? 5 ) else ha- a fonsls set cf uro¬ 
genital slim© d frivstlve •. fe wive is open sid *, the bulbe*ve ti hilar 
^Unds' are sell developed, th? vsc^ibuls ;ide and rhnrt* the ubursihral 
divert!cuius: large, Wolffian duotr, Join the vagina, tfe f :s both Join the 
veatibuls, and fh-* suurrOwed itr thru 4*l/*i*4 ae Ion as in th© fossale 



59 . 


of this side while the cmle has it l.ff/^,4 as long as the finals* the 
a'all prir sordine* of th? P- ra-urctbral duets ap oar an in a fessale of 
20.0 or* (?ir. 11) on the dcrac-lai ral groove o vssiibule cu a*iCh 
side. Tf this i & horologno of the pro state as givan by eu. sbaaar 
C 1 ** develop* 3-stc? here as in t Mai#* and ir the female fashion, 
not pestaro-antcrlorly as in the sale. 

The 20.0 Ofl. frea-aartin (fig. £4) has & such acre advanced fe- 
r/sale typo of vulva and a flattened cut v lvo-vestibtiiar canal, eh lie 
the bulbo-vestibular glands arc better developed. Cifcer s^v^nceu in 
devolo-,sent u e e own by the r ortening »"d widening of the vestibule* 
the ne^.hod of union • f the vagina auaoulina ?rith the oiffian ducts 
and sinus, 'the suburethral diverticulum, the length f the urethra 
pr psr, and overs ir th presence and condition of th.- two nil para¬ 
urethral ducts -f Skeane as in the 5 .0 on. fesaio, but larger. And 
so hero, too, th‘ female conditions are retained, but tbu* para-ure¬ 
thral ducts nr ahead of the fenale, but are evidently « eendary in 
their tine o atirrulation, Just &» in the ferule, but thia study dc©3 
not tabs in late enough stages to decide their ex id hoaclogy. 

lbs sexual differences in the develop® at of the derivatives of 
the urogenital sinus and plate, brought out by i i study, are a* 
follows*- 

(1) The devalopaant of the lon t , cavornocs urethra with sigaoid 
flexure in the sale, lacking in the ferule, ic aocosp&aisd i . the 
ffiiile by widening of the psrincun a d closing over of th urc ratal 
piste, with consequent vontro-artsrior olgration of the penis. Its 
aba once ic the fa ale in aoc epanied by greater recurving cf th© 
clitoris. 

(2) Tovolcpaent of the avail supra-urethral divertieolw* in the 
male at tho bend of the urethra and junction of + h cavernous and pro- 
static portions, to receive the bulbo-urethral gland*, with no re-o'ccu© 
in the fera! i. 






































' 



















































«Q. 


(3) The long narrow prostctio portion of the urethra i-*. the sals 
as coot rat; tod with the wide short vstibule i the female. 

(4) The difference ir. type of the bulto-arathral glands in nalee 
fre* that tf the bulbo-veatibul nr * in fb© feaale, their hosologuss. 

(&) The development in the sale of profuse prestaios, eurrouniing 
th© prc-.tatic urethra from the fcolho-urethral gland region to the open- 
irua of the folffian iucte, and spreading in a pentero-anterior direc¬ 
tion* they &p mr firnt at 5.1 or-. to, . The para-urethral duct#? of 
ti»e female, givon fey ^©ugebauor a ♦be horolcguoit of the prostates# do 
net appear in fecal os before th t P 0.0 os. stage, when two or throe 
<*p ear in the dorso-lati/ral groovaa of th vestibule, near the vagina 
Opening, not near the bulbo-vealibular gl&ndn an in -ales. 

(6) The absence in th • n&le of & hocolo/ue for the sub-urethral 
diverticulum of the ftormle, and the failure of its urethra to narrow 
in fr :nt of the folffian luot openings a - in females. 

(?) The short urethra proper 1 r the male as contrasted with one 
twice as Ion or sore# in the fe*ale. 

(8) The great development of the long corpus cavernoue urethrae 

in the a,la# with it: aignoid flexure# a- coatr-cted with its alrost 

* 

rudi?>ontary condiiior as the corpu c&varaosuR clitoridii: of the fe¬ 
male. abort and ainocat. 

(9) The opening of the Wolf ien ducts separately anterior to the 
vagina masculine in th? ta 1©, ac contrasted with their union with the 
vagina before opening &s one to the vo tibule in the fir ala. 

In all thi? above features, er.oopt th© last, th© freo»nertir» is 
definitely and ««ai©tafcably female as far as shown by the specimens 
studied. he etructcree mentioned in (9) above are of aoeoderml 
origin, which say in a aeaour© account for their being influenced in 
the sale direction, while the rest o* th© alaufi derivatives, oac *pt 
the strip between the ureters and the olffian ducts oo iiiu uorsal 






































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• 










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61 . 


urethral vail, are of efttodermal origin and are essentially unaf- 

y 

fected, and remain female in mode of development. 

The Phallus and External Genitalia. 

In females: 

The female external genitalia differ from those of the male 
in the perforation of the urogenital plate in early stages. This 
takes plaoe before the 2.8 cm. stage. At that time the position 
of the phallus is approximately the same in the two sexes, and both 
have genital swellings and teat primordia. In the female, as time 
goes on, the phallus becomes more recurved toward the anal opening, 
the vulva more open, and the perineum no wider relatively. The 
genital swellings in cattle, unlike those in the human, as described 
by McMurrick in his * Development of the Human Body", do not take 
part in the formation of the labia majora, as is shown by F. R. Lillie 
in his unpublished work on the homologies and fate of the external 
genitalia in oattle, but completely disappear in females at about 
10.0 cm. - 12.0 cm., and labia majors are absent in cattle. The 
lips of the genital plate form the genital folds and, finally, the 
labia minora. The plaoe of the absent labia majors is taken, in 
later stages, by the labia minora or nymphae, which grow out on the 
sides of the recurved phallus and approach its tip, finally over- 
lapping it in the older stages, though not by the 20.0 cm. stage. 

The tip of the phallus is cavernous and forms the glans clitoridis. 

In early stages the glans has an undercut groove in its dorsal part, 
filled with stratified epithelium. This disappears in later stages. 

The thick ventral wall of the vulva becomes cavernous, as the corpus 
caver no sum clitoridis, which is sinuous and does not have the great 
increase in lenght so characteristic of the male. The glans clito— 
ridis is knob-shaped at first, but by the 20.0 cm. stage it is pointed. 
The udder and teats develop and the genital swellings give rise to 




* 




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' 

... • * . * 

. 

- 

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. ’ - . ■ ■ • 

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. 

• • ' : 

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- . ■ .. . - • . 

' ' ' - ' 

.. . 

. 


sxa&Il scrotal prominences which disappear after tha 10*0 ea* stage. 

In Balers: 

Hie phallus derates along the ventral body wall, following 
failure of the urogenital plate to perforate, as it does in females; 

4 

the perineum grows ventraily over the dorsal part of the plate; the 
lips of the plate ft$se to produce a raphe on the body wall, thus 
the phallus is pushed forward to the unbllicus region, and ooa^s 
finally to point antero-ventrally. .be sigmoid flexure of the much 
elongated penis, including the urethra and corpus oav^rnosun, develops 
fro® before the 7.5 css. stage (Fig. W), The genital swellings 
develop into a scrotum, no udder develops, and teats rerain rudi- 
aeniary. The sides of the phallus develop into a sheath of skis 
aa a prepuce at about 10.0 oa« (Fig. 17 ). and a fe sa develops round 
the glass penis, which glam? remains projecting forward as the end 
of the corpus c&vernooun urethrae. The glano remains open ventrally 
allowing the urethra to op*n on its ventral face, and is attached 

w 

to the prepuce by a frenulum or atrip whiob takes a spiral course 
along one side of the ^Zans, from a ventral position near the tip, 
to one slightly to the left at the botto© of the fossa. 

In the free-eartin: 

The phallus development of the free-mrttn is completely 
and exclusively cf the female type, so far as seen in the 36 free- 
sartins collected whil^ pursuing this study, and lillle (*17, , S.-) 
decs not record any modification of the external genitalia in the 
sale direction. Keller and Taadler (*ld) oention finding aoao 
such modifications of the external genitalia, but do not describe 
them. The only oaaesproperly d escribed, d« that of t ha—n (*43,'44), 



Tuoh esses occur in single born calves, and therefore need not 
be due to the situation during intra-uterine life, but nay very 
well be sygotie intersexes, of which one would expect to find 
the sane percentage among frse-smrtins as aaong single born heifers. 










# 



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. 












. 

| m - . • J ^ : i 

« J • ' • : . j ■ 






' ’ < 





65 * 


and of which ther* are so e« recorded cones. .hero is sous reduction 

/ 

in di&e of the external genitalia in some oases* e. g. the 9. ) cm. 
freo-tsartin $ £• But tier is no eign in any of these free-oartins 
of forward migration of the gten—« nor of olosuro of tbs vulva by 
©vtsrgrowth of tb® perineus as in the sals* 'her,® minor redifieationa 
found ars all to fee found in normal f etudes so far as acted * in 

f 

freo-esortins the scrotal prosdoenoes disappear as in the normal 
female, and the «-;d«r forss, but atill remains snail* even shoe corr# 
latod in the individual with descent of the gonads as in sales, 

l-acaary and Diacu sic s. 

(l) the ^on&d of the Area■ ■nrtio dswlope as an ovary to 
a varying degree of di ff erentiation * depending on the tisKO ©f 
initial reaction to the sals hormone, and say e co fur is the 
fesale direction of ite differentiation aa to develop a considerable 
layer of secondary aex-cor s, w.ich* un.er the influence of the 
sale borsenc, undergo degeneration* and apj^aruntly entirely disappear 
‘,vhe sex-cord region* in all cases exa&ined# is cut off f^ox the 
epithsliur covering the gonad by a tunica albuginea resembling that 
of a Bale of smaller si so than the free-martin* and the ret® of 
the gonad a &usr.e a position in the gonad correlated with the degree 
of development in the sale direction reached by the other acx 
organs. 

(S) A sexual dif ference is evident in the such 1eager 
distance on the urethra b tween the openings of the ureters and 
those of the lolffian ducts in tbs female as compared with the 
i&ale* these structures have a cosiaac origin in both sales und 
feoales* but the ureters migrate anteriorly along the urethra in 
both sexes. Ihe free-sarUn is usually intermediate in length of 
this tube* but is eoaetises oven longer than the corresponding 
female. It does not approach the sale coalition in the oases 


studied. 





64 . 


(3) In females progressive narrowing and degeneration of 
the Wolffian ducts set in at about 5*1 ca* stage and continues* 
at a rate varying in different individual** till* at 15.0 crs., the 
ducts ar- intere&ttant ant rior to the vagina* and, at 20.0 cn. 
they are present only in the uterus wall, as solid strips* and as 
sdnute isolatad segaents, in the wall of the anterior half of the 
▼agina* but continuous in the posterior half* and with a lun?n in 
th* posterior one-third of the vagina wall. They have lost all 
connection to the rate of the gonad and to the cranial group of 
meeonsphric tubules. 

In sales these ducts continue to develop and have a lixnsn 
throughout. Ihey say even occupy the whole of the genital cord 
in places shore the yallerian ducts are degenerated conplotely. 
They retain a connection through the cranial group of wosonepfcrlc 
tubulsa with the rote testis, in the epidldysds. 

In froe-aartins the condition of the ducts varies. In one 
case (11.75 era.) they have degenerated in the anterior part as in 
the femle. and have lost all connection through the cranial group 
of mesonephric tubules to the rote. In the other cases studied, 
they ratal this* connection and are r*le in condition, but are 
comparable to stales of younger stages. In ail* they appear to 
hava behaved as ia the f aal® for a considerable ti*;, and to have 
degenerated somewhat, and then been stimulated again. 

(4) Ia females the Wolffian ducts in later stages Join the 
vagina before opening into the sinus, while in males they open 
separately throughout the series as in early stages of both sexes. 
In frve-aertins the condition is variable. fonetines it is as 

in fosales. and at other tinea as in males. This bears a relation 
to the degree of maleness: of the other organs in each individual. 

(5) The sesinal vesicle priaerdia begin to show development 
in the 7.5 o». females, and five, or possibly no re, begin develop- 
sent in the stages studied. But non® reach a else appreciable in 



gross discretion, and in later stages they appear on an inter¬ 
rupted Wolffian duct* and* in rany cases, on isolated parts of it, 
and always farther forward than in mice of like aise. I the 
laat stage studied they are found in the lateral wa lfi of the 
cervix uteri and anterior part of the vagina. The number o* minute 
vesicles is a sex oh raoterietic. 

Tft stolon usually one, or at ~o*t two, occur, of which the 
posterior one dooc not develop, while the anterior one on each 
aide grows strongly to a considerable length before branching and 
the opening of its single duct in later stages is progressively 
nearer the posterior end of the Tol^fiao duct, into wh eh it crens« 

The large also and preface branching of the single seminal vesicle 
on each side, in the male, is & striking characteristic of the sex. 

In free-^ariins the condition as regards s?Rinal vesicles 
ie varied. All appear to have developed as females with coae inhibi¬ 
tion of the vesicle growth, followed by increased stimulation, 
which causes increased else, like that in the sale, but with increased 
number as in the female, or even more, e, g.,tha gO,D cm. fres- 
aartin with 11 diverticula on each side. This is again a case of 
a male typo of development, imposed upon & sot of female pricordia, 
the degree of which &pp ara to be a function cf the time of incep¬ 
tion of the mis feormene, a&i the time elapsed from inception to 
the stage studied• 

($) There appears tc fee a sexual difference in the rate of 
growth of the Pierian ducts in stages that appear to be sexually 
Indifferent externally. In females the Hfcllerian ducts re ch tho 
sinus earlier time in males, and as this occurs in tho female, at 
least, before either sex has interstitial cells, it is believed to 
be xygotioally controlled. 

(?) In females the *s&llorian ducts centime tc develop into 
the cstium, fallopian tubes, cornua uteri, corpus uteri, cervix start, 
and vagina, and reaain continuous and patent throughout. The 
retrbat of the gonad and the altered proportions of the pelvic 


























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9 : ^ ^ is • si ?-*: ^ •' • * 

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66 


ration oaune a looping of the cornua irt&ri and tubas, The vagina 
and the utsruo increase greatly in diaaeiar in later stacec, and 
the forsor opens straight posteriorly into the dorsal corridor of 
the vestibule. 

In mleo tho ostium degenerate* and leave.: only the Jiydatid of 
SftOrgii-nl, which is no* always present, oven up to £0.0 on. itagea* and 
the duct itself degereraiec completely in the tubar and co~nua uteri 
regions. In all th? stages studied, a r^sami of the corpus uteri is 
found as ^ vesicle, blind both anteriorly an* posteriorly, at the 
junction of *fe» genital cords, or a~. a solid stri in the region wh- j r« 
corpus and o rvix uteri would be in a forale. The vagina rascalina 
is prsoent in stag s studied, but hug degen rated to a vesicle filled 
with epithelium, dorsal to the openings of the olffian ducts into 
the sinus, by the £6.0 O*. rta?o. 

In frce-mrtinc d generation of the ^llerian ducts gees on as 
in the sale, but it hae started later, a^tar the duct h*d attained a 
more advanced state than in the norm! sale, and, w? il degeneration 
seems to h&v# been proceeding sore or css uniformly throughout the 
duct length, it disappears first n ar the anterior end, in the tubar 
region, and la t, whf»ro the utertm and vagina wo** Id bo. Here again 
the two 9.0 crs. free-martins show e variation in th-> condition of 
these ducts, correlated with that of the gonad and *©lffian ducts, 
which points to a difference in time of onset of the wale herraon?, and 
consequent 1 n&er tine of degeneration in Ho. 2, acting on a less ad- 
vanoed stag* of female differentiation, an oonpared with to. 1, with 
later onset and rhorter duration, acting on a ranch core advanced 
stage of female development. In the oldest freo-^artin no hydatid 
i* fo* nd 03 in com ml?s. The other three present a series in do- 
generation of he duct derivatives as in males, except that the onset 
of degenerafct on ha occurred at various stages, and therefore has 
reached various stages, ^Ivayo cosparst-ls to tf^ose of younger miss. 

(8) In both sexes, and In the free-nartin, the state of do- 




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6 ^. 


velopmeat of both sets of duo*s on the loft aide advanced 

than tho« e on tho ri.ht sido, arid thlo is particularly true of ievelpp- 
aont in tbs sale direction. The retreat of the rcca ana of the . %.z&- 
nephro: and the loop of the ge-ital cords all ; ivo evil not of this, 

(2) In fersalss, the ancon* vaginalis usually fails to d vs;lop 
to any conidurable dopth, while, In the sals, it gresra vvntro-laiertlly 
through the Inguinal canal to th * top o* the *croturj, in the specimens 
studied. The deriv&tivra of the ingtr!no.1 IV sent, the round lig sent 
of tbs fcs&Ie, and the gubet'anculur fcrsiis, in the wale# tfeile they 
behave in such the sane amwr, a a nade to appear to do differently, 
by the differing development of the vaginal process d :ri?ativos, in 
the two sexes. oth igaa nts elongate, then thicken for a tire, 
then shorten; and, in the ;tjue*> beyond tbi? study, beco e reduced 
to their definitive condition, the tub© racul tats in th fcoft'-s of the 
3crottna, and the round li^aaent in th© body cavity* Th the stages 
covered, by this stody, the ronad liganaat lengthens and thickens In 
the abdoaan, fcecan :e there Is co aaccus to receive it, and the guber- 
naoulaet lengthens, partly in the abdeeon, but mostly in the succus 
vaginal!.. , t.dj's.i thickens raolly in the ;? siccus* F*art of tho ro ,-ri Xi£&.~ 
sent is taken up into tbs sails of the cornua uteri cl o, but it 
thick; ns ali.hlly ?coro than the guborncultm in the stages studied. 

Freo-sartins ars inb-raediai©. the ease diff ranee in decree 
of stil^nses is ouad in the Vo fr ns-raartins of tho s a age (3,0 ©a*). 
The others give a series of stagea in which tho ©scows is deeper than 
in feaalee, but <3i co deep as In svxles of tho cars age, ani the 
thickacing of the inguinal ligaaent derivative occurs, partly in in* 
saccu 2 , and portly in tho ohicNn* depending on the depth of -be aaccus. 
In the. two latvv.it eleffj f ta tf Ml for this paper, the WHI had pene¬ 
trated the inguinal er.aal c-n bet but in Mm cases it penetrates 

on ono side only, end that side develops as in a oaie, while the other 
sido develops ac ir: a feiaalo* .here ro Id *i>p*ar to be a tis© after 
















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1 

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6 ». 


which the canal cannot be penetrated, the left side coccus soas- 
tir^s is far enough in advance of the right one to penetrate* while 
the ri.ht does not* In that case the left aid* develops a typical 
guber r:ac ilu the rl«fet, a typical round li luaent. 

(l0) There is a sexual difference i the netted aci degree of 
io. en ration cf the steaonephros* “ ; esit takes place ouch more 

uniformly throughout its entire length» and the cranial group of tubuio. 
do not retain a connection with the rete of the gonad an i the olffimn 
duot, an is the case with sales* but the rete of the ovary retains 
•case connection throu h the degenerated a ■sonc-phres to the ostins* 

An epoophorors, tram. the cranial group of tubules, and a paroophoron, 
of a few tubules at tha posterior end of the gonad, frois the? caudal 
group of tubules, reaain at the 80*0 cm, al& o. 

In rsales, the cranial group o? tubules retain their connection 
with both rate testis sad olffian duct as p:rt of th opi idymis, 
and do not degenerate as in females. he caudal group of tubules 
degen rate?? as n the female, but raaaast* forss the paradidymis, and 
a single inol&ied one, the vac or ducto’ sberrans* * Ihe whole neao- 
neohron is slightly slower to degenerate in the sale* ana th:* rote 
testis doe? not connect to the ©'tier* looerali dieapps&r af ter 
1?.0 oa* ata ; e, and before £C.Q cm. stage in both sexes* 

In free-’sartine, in all those studied, except the 11*75 on* 
one, where the Wolffian ducts are absent in the anterior region, the 
era ial arotip of tubules act as u connection ot th- olffian duct tc 
the rate of the gonad m in sales t and, even in that cate, with no 
ant *r or end cf the Wolffian duct, the cranial fere up of tubules; le 
retained. In ihi later stages it far as part of tbe epididymis as in 
miss, but i:; behind the normal mal^, in state of development. In 
early stag -e studied, th^ rate is connected to thi omtiu remnant, 
though not so wall an in females, eh le in later stages it is not, 
and res rubles a younger roils condition* The ohtPWt^riitio paredidynis 


/ 



tt(e -h 

70. 


and vas aberrant , o-" the ml.-* ar- found in tho later ata as, with 
rsore woo one pj.ro s: remnants than 1 either oox. The free n&rtin follows 
the Sitie in deg ~n ration of the nssonsphros, not the female, but some 
si na of the early female condition are present bora also. 

(11) fbo development of the urogenital sinus d rivolives in tho 
fsisalo, sii>lo and free-wartin, in “utmarised in the d toiled description 
of those organs ar,d their development, and will no? be repeated hers, 
except to point out Of in that in fres-iaarti no tho characterization 

, iven for the female is eq ally applic&blo, with the exception that 
tho anaer of oponirr of tho vagina and ’olffiaa ducts int? tho si nos 
varies; all the fro^ artins CM nho M os. So* 8 specimen wore 
fe.aa-o in that respect, and this one, in otho re..pcois, wao in a noro 
advanooo sals condition than the others. Those ducts are aeacderrml 
in origin* and the r-^st of tho structures iarlvsc ■ the sinus • 
tirely, are entoderm* in origin. 

(12) The normal o^v«lo w&n t of the rale and ferale external 
genitalia are a scribed under that heaiinj. in the general descrip¬ 
tions, a was also that of tho free-wartin. It will he worthy of 
note here that there was not found in any of th« froe-nariinr used 
in this study, nor in any o the 31 others collected whil* pursuing 
ti.is investigation, any sin of modification of ext r al genitalia 
in the wale direction, there is some tendency to smaller Rise of 
ex ernal genitalia in sobs Of the cases, but th.y a », in all cases 
exaniaoci, completely feiaU* * and hawe otiiy such minor variations a 
are feuej i- single— f?orn fe®al s* he .cro*u fails to forts, even 
shea de cent of the gonad? talc s place, a in rales, and aiders are 
pr u’s ?nt in all eai>£:? examined. 

( 13 ) The d*ta given above support th© theory of hillie that 
the free-sartin condition is tb result of the action of a ale sox- 
hornone♦ probably eecreto- in the inter titial cells o the testis 













































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r 









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T! 


of the ©die <o»6 paswuid to the ay, .ctic*lly dstenda«4 fosaie 

iu ut«ro by pi«0 at&i vaaouxar armstoasoai-.: , ana that this fecrsoa© 
modifies Use f^a*© is the aal» oirootioa mijalo the femle a*, oo 
influence cm the »-ie, 

(--*) It is fcalievea that tho factors una rlyin^ the different 
ef f ots am cegreea c tiio sue*> effect of the heraose in 0 Iffereni 
frem-curtxna of th s«*i^o aise, ash. in t;.o»e of different tiees Wi»- 

(a) tim* ©.: initial herwoe action, which ia tiara i* dep&xfcient oo. the 
ti*e of horaono mo oration by toe sale co-t»i, or or. the blsae of fonaa- 
tion or t-h pi c utei vascular ^.n*a toaosie, 

(b) the length of tii?e the i or cue ha been active# «;deh taken with 
the oonoitior? at t mo ©f onset, loads to the elate arrived at in any 
stage, as in imiea, hit - . th the cifferanoe* that i sales it basins 
to act tm a conoii on that 1 approximately coon taut# efeile in tfe®- 
fr»«*Ofe rtin the i. :,itia. condition i ou.njeot to variation d i-pendent 
on (a) am probably aln© © diff^reooos i *y. ctic^lly controller 
diff rent iat on of tb organ# at Um © on at, &r u,. eatocl by illie 
f f 23) and afcom in a eemnnris f -« of tm> lb#-j oa« and 16,b on, normal 
fe* alee describes In thta stony, to fee to* eu. e in later atages, 

h<- ■■ z v o* ic difthraaeai My ol o ntftort tfcft ftflrt P I VtSo Bent . n.: r 
aaSc-boroore influonao, »** deter^dLaiiie: the final condition ar?ivoa at 
in any specimen studied, 

(lb to all cases ©ba rred# the effects of thi borrnone i t©«* 
rale direction have been li tad in organa of a sod rnnl origin, and 
do not appear in those of (otso^md origin oxoept am *-©taa for para¬ 
urethral do t», 

(Id) As to the ™o 1?' of action of the hornets , tr iyi»® of view 
may be b«ld;- 

(a) tnat it is ©pcci ic in its action «*d affects each organ or s*t ft 
organa iadof-endnutly* by it? pccuiar ohesicai conatitution, in a defi¬ 
nite and purely qualitative easier, than eaucing purely qualitative 






























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72 . 


charges in each organ under its influence, This* places the specificity 
in the hcriso e* 

(h) th ti* it h„: a lu&nUtiitive v.ffect rather than * 3«al: * stive one, 

\ 

a*v< .iter th© t . emral life proosse a opaniitati^ely, Thi -* ay ; ?rhap« 
ha ‘ ht of £e& fectin the rate o heael i ft*, bolls-, Qg i« ly rais¬ 
in...; that of the forsale up to, or nearly op te# that of the s&lc, thus 
^llo. ihy each orgau or set o r organs to re-act in it' ©peel io way to 
th^ cha. gi>d conditions, end ir that d■.*free which in celled *orth fry the 
pa: 4 leaJar m-tafeolic ryt reached in the individual, and correlated with 
the different rations of the tissue itself, This places the sp cificity 
in the titan^a tho^seiv s and in their inherent ty ? o reaction, and 
cons:drs the actic of the borrore as purely qu ntitativ , 

fh .e it in not believed that trig a 4 tidy in any ©arise settles the 
question as to which is th© nod© her * at work, 9 *>®e of he f rdio 9 are 
8$2£g>. live* H»r*y are a-, t noble © r irt■*rprotation either way. Th© 4 a-- 
generation of th© secondary seu-cord* or th ir n n-app-arauce in r:my 
caaer. aay b^ xpla nod either as due to an altered general physiologic 
0*1 react on w lob is inhibitory to its f. rssatio acri oauees its de- 

• 

generation, or else the horrsona aay fee thou ht to be specific in astir 
dir sotly on th© tie ue. 

vho r^t‘i:n of the ilfclleriiw duct say be a specific reaction to 
an altered rotabol c r te, cr the hormone may be specific i it# action 
co thi d :rivaiiv<js of he duct. If the latter view la held, it Kay be 
ashed, why the different parts of the duct which are derived from the 
same tissue, and pr^su-ably receive the sane cose of the hormone 
throughout in one individual, nei her develop nor degenerate at the 
aune rate throughout, hut my be pr «nt in a fair ntate of pre¬ 
servation in one place and entirely gone in others, both regions 
having a c zar.on origin and th» e*«S 9 dose of hor* one. This can perhaps 
be easier understood be expected, if the horaone in ot looked 

upon as aboolut:ly apecifi© in its action. 














, 
















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' 





73 


ifce Wolff i&a. duct derivelives, particularly the s«ai<ial veaiels*, 
in tbeir de?slop:3:nt, in the iw a «ni ic the fro -ru^rtin, cay be 


lEora eerily explained on a basic of quantitative reaction than on the 
purely qualitative on** -he developn-nt of one, „ni at soot two seal- 
nal Ys.'icleo in nor:: 1 males# of srhiob only the nt ri-.-r one develops 
tc any tq^reciahlc ai-.a, uad t ho appearance o: four or five i: tun fa- 
' olOi of which aone grows strongly oaough to bo uD'ioe&blo in gross 
dissection, con be easily explained on th: basis of a specific horaoas, 
:±r.d *hs l&ck of it* or on tfca basis of an altered quantitative physio¬ 
logical reaction in the organise* In the froe-aai*tin, the growth of 
■ 3 1 rvon o these trticula on oatfh ids, aa found in end c&oj. to¬ 
gether with the evident nal: type of davolopnant found in aeveral of 
then, out to a 1 as degree than in the isale, and the evident graded 
aeries that they forr., hardly look like a specific korsone action, 
pickin'.; out the Sale structures for stimulation and inhibiting the 
ferale ones, for it wo Id stimulate only the ant<vior one; bit can be 
under tood better, perhaps, on the basis of the development of C-aaon 
pri ciordia under one phsyiolOoioai coriiitioc up to a certain state# 
then stimulated as a whole t a quantih& lively d ffereni condition, 
ifee varying degree of development in the sjpI .s eleven vehicles ia 


a u&ntitativ' d fferenoe, net one of quality. thi3 is very sauefc like 
the iort of result produced A pie’ reaction ay teas by w dely dif¬ 
ferent choice 1 re gents, which work principally by altering th: 
hyaiological rsaotion quantitatively. 

.he earlier appearance of para-urethral duct a of ferale tyre is 
perhape hardly the action of a ©I’ecHio rials borvone, w- icb should 
pr oduce pro. tales of scale typs. not neroly crtiaul&t* the pri^aordiu® 
of tho female structure. 

7h relative lengths of the urethra proper and the typs and de¬ 
gree of degeneration cf the saeeoneptjrds, are capable of explanation 
either ray* :fce fact that that of the free-awrtin usually fallows the 
sale, but i« slower than either sex nomxU is a quantitative difference 



























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74 


ac veil as a qualitative ona in any case. 

Ths failure of th urogonitai inua derivatives generally, to be 
af'®o*!id ir exp intable either way. It uny b? due* either to lock of 
sunoepiibility t altered phy ological activity, sr to lack cf sus¬ 
ceptibility to the action of the specific horccne, possibly because 
pant the critical sia&» before the hormone b ;cttsc effective, in w Ich 
earlier introduction -i ht affect the , or to feein coapletely 
under sy otic control. 

vjen^rai changes in body idiupe and bah >vior betwe-.n tno two soxjb 
and frad-aiartin, while it i c...pable of satisfactory explanation on the 
basis of t\io qualitative specific iior»; no, io equally sc, on the bards 
HP a quantitative change i * ph>biological rreeti os initiated by the 
eey: hereto no, and work! g on tissues throughout the body, which react 
•pocifically t it in varying degree, depending on their own physio¬ 
logical conditions r lutiva tc it. 

(17) That the her one rust b© continuously passed to the frse- 
rariin to sustain the development or d*£«n ration, which it initiate?, 
would appear >012 tfc;> offoct of castration on th* general body ah pe 
of ‘he me' if 

aft 5 r castration a-, shewn for guin©a pi e by faB'nis (*0' ), Fisher (* ’3). 
They do no 1 null] i cattle feral es, and are pre uhiy . .v.ondent < 
the tcartic in rml and it would appear that It i expectir:. such to 
suppose that the ef ect on these and other atruetores in the free-jaartin 
is not dependent o« the tire the- hormone haa beer active cr that if the 
horaone has been active oni has initiated the development of the free- 
martin condition, that condition will develop to completion, if the 
bornone i* cut off before aucii completion it* arrived at, ev:n thou b h 
ite. threshold o " action ic low, ana, wnile present, it appears to follow 

the "all or none" law in action aa sug<eatea by billie (*21). 

. 










































. 
































■ 







































3.iter * ro cited. 


basc-ors, •'« 1., 19**3, ho Interstitial Jails of tha .enada of Cattle 

with cpocial iefsre no® t: * h ir ebryonic 
uovs lopac'ct one, ;i r nificunoe. 

An. Jour, of Amt., 31, 1923. 

at ms or, 1913# Frobleee o* r» ties. (pp. 44- t) 

Idle l? vor - ity Presn. 

.er solicitor, Aa•, 19 2, fie *11 .rnchen and olffocfeer Eru.e und die 

dildun. des w~iblichen . onitoltractus helm find. 
Arch. Anal. inu Phyc. Amt,, Abth., pp,2-.. . • 
Jhspir.# Catherine#!,. ,19'’ 7, a -icrc icopic f? udy o r the productive 

System In foetal Free—nar* no. 

Jour. ~xpt. loci., 23. pp. 4^1-482. 

Arch. Di Ana to si a, 7, p. 294. 

?ho Development of the Urogenital Organs . 

!anual of •iu©»n frbryolo y, Koifcoi and Mall, 
V# I!, pp. 252-979. 

fho Influence of the Jonai Boraonss on the 
Seminal Vesicle. 

Amor. J r. of Phys., 64. 

John ii' ntor’s bree-eartin with ^eoarrs on feme 
r ecerst Irvo a* Ig® 41 or n . 

' Unbar, l «d* Jour., 20, pp. -4-93, 

The Freemartin • it’ C' - •. c - ph‘-nr, 
dan, < o... • 

C . . a l 1 * a 

roller# X. and Tandier, J. 

1916, debar do ferhalten dor ih&uto bei er iwillur^ 

* otrachtigkeii d?s indoc. ntersuchun&on ueber 

die nt• toftun^sur n ache dor , o chlechtllchen 

Jntorentwioklur.f von ns*. r 1 icfc s ''.williu. rskelborn, 

. « 

seiche einon -k* ni ich .-r. albc sur ' nt*icfcolti*\. 


.✓©ftonis, 1908 , 

Felix# 191 ?, 

i..her, I, * 1926, 

. f»rt, .) • 3arry, 192 v • 

hariman. Car;. .., If70, 
























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Literature Ci ted. (eenst *d) 


Lillie* r. R., 1?16, 


in 7 . 


1023, 


Lillie, i, a, and 

fiascos, i, 1992. 


gelurgen. s*ner ':iar&rs.tl. 

seer onechrift,, III Jabe*. pp. Ll3-.2$. 
the beory of the Fre-eartir** 

Science, 4v» 611—613. 

fhe Free-wertint & Study of th motion of lax 
Hcrooiseu in the F©«tal Life of Settle* 

Jour* Kz . iooi., £2, 371-it;• 

Supplea**' Lory Uoioi, on *wiaa in Jattle. 

Biol* Sail** 44* 47- a. 

an Lariy 3t%9 of the Fr c-aajrtia and the 
rartlel victory of *ho Interstitial Jells* 
licienee* ii* S*. be, 143£. 


J&ox-, 1913, cited fros Lillie, 1923. 

sui^nucacr** h., 191 «, oticnlohtslosc L« Ullage, Line *J*Shnli©fce 

fOns ecu uerxaepnroiitianus beio Einde* 

irci*. fur. _i.net. unJ Ffcye. clogis, s&at* 
libt* 1 * Jj 1 —Jij* 


Jo.'lurrlok. . * 


»» 


ilooro» ->* , 1021, 


Ihe Develejpaent of the hueen .©ay. fifth d. 
Ob the Phyoioic„;ic*I Properties of th« omdu 
as CSoistr oiler.. of ©sail© «*..© fayohical 
Characteristics* 

Sit* artificial herstt^hrediti* in fats, 
jour. ixp, fool., 33* 1-61* 

Strsan, 1S44, cited fro* Lillie* 1017. <- 

fend, K ud, 1917, fixperlu ntn on the Internal Sucretion of the 

3 suai 2lands. J&pcoially on i.xpe io^ntal 
iiersatphrod itiea* 

Jcmr• of Phyoioi.* 53, 207-063* 


*;p Igalberf, Otto, 1361, Uebsr die rktaseruav; her ..enitfclicn bei 

(en&eblick) v©raebii>a n geaefleehtlichen 
Zwillie&skhlbern* 

Zeitcchr. f* rat. ed.* Til Reihe, 3*11,120-151. 









* 


















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♦♦ 

























Literature citod (cont’d) 


Stoinach, 1020, Kuntsliche tmd nat&rlioh Zwittordrfrson und ihre 

analogon Wirkungan, 

Archiv, f* Snt*r.-!?lenh., 8d. 46, S* 1S-56. 

Tandlor, Julius and 

£ell«r, Karl, 1911, Haber dan Varhalteis dea Chorions bei verschiodjan- 

geschleohtlicher Zwiilinpegrarriditat daa Eindes 
und fiber di ? ^eroholo ie dea Jenitales der woi- 
Mlch'-'i fieri? seiche ainer aolchsn ;rariditAttnt- 
8tUT3«n, 

lautaoho Tler4r2t. ocbenschrift., Janr... 11. 

Miliar, B. H,, 1920* Structures and ?joaolo^i»s of Free-nartin onads. 

Jour. of Ftp. gooi., 53* 65-127. 
































































































































GLOSvlORr OF HOUOLOuOU: STTJCrJRSS !* UAL2S, F^aLSG a:J 0 Pfc2B-MAm3S. 

Appendix testis* Hydatid of Morgagni. or Sessile hydatid;- the remnant 
of the fi&bri&e of the octiuxs found in iaal?3 sod froe-rsartina. 
(leugebauer. ’curriok, Pre tiss and Aroy.) 

Bu bo-urethral or Oowpor's glands; hoaologuea of Bulbo-vsotibular, fculfco- 
vulvar or Bartholin s glands in females and free-nsartinc; - compound 
tubular glands open!ng into the sd nus between prostatic and cavern¬ 
ous parte of the urethra. 

(3ougebauer» Prentiss and Arcy.) 

e 

Cavernous urethra; homolojue of bride pniann® or ridge on the under sur¬ 
face of tho clitoris in fesales and frec-martin 3 ;- the part of the 
urethra adjacent to the corpus caverr.osuE p:nia or clitoridis. 
(Seugebau r.) 

Cervix uteri of the fenaie is that part of the fused dbllerian ducts 

whore the constriction occurs between the corpus uteri and vagina. 

descent of the gonad* and Mesonephros;- in both sexas the migration of 
those organs posteriorly as described for aan by citurriok. 

Luctuli eff©rentes of the cpididyais; houologu » of the opopfaoron or 

par ovariuc or organ of l ooenrtbller:- the crania group of ueoone- 
phric tubuios or thuir remains, the rote testis or reto ovarii. 
(ucUurrick) 

Cuotus epididymis. Vas deferens of sales or free-varfins;- th^ homologues 
of the canals of the epcophoron or artner’s ducts o p females - the 
derivative of the Wolffian or i4esoncphric ducts. 

(Prentiss and Arsy) 

Jenital cord;- the ridgo that contains the olffi n and ^hlleri&n ducts 
froa the front of the aeaonephros to too uterus region. 

Germinal epithelium (both sexes) those layers of 4 he early ^onad covering 
that continue to proliferate inward to give the sex-cord? and the 
antecod-nts of “ho gar -cells. 

(Willier) 


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2 


Gl&gs penis; homolgoue of glans clitoridis of females and free-rartins - 
tip of the phallus cut off from the prepuce by the coronary sulcus 

and attached along one aide and the definitive ventral surface by 

✓ 

the frenulum of the prepuce, Rudimentary in fenales and free-martins. 
(Heugeba er, Prentiss and Aray, ? o’Jurriok) 

Gubernaoulun testis; honologue to the round ligament in females and termed 
\aa eith r according an it develops in the saocus vaginalis or in 
the abdominal cavity in froe-martins, - the ligament that attaches* 
the genital cord to the bottom of the saccus vaginalis or such 
homologous structure as may bo present derived frcm the vaginal 
process or canal of Buck. 

(Mclhirriok) 

Medulla of 'he gonad;- that portion of the gonad forced by the primary 

* 

sex-cords destined to degenerate. 

(Sillier) 

Kembranous urethra;-that part of th-i urethra nearest the vulva in the 
female ana freo-raartinjhomologuQ3 to the part of the urethra 
between the prostatic and cavernous parts. 

Ostium tubae abdordnale of tho female i3 the open anterior end o f the 
oviduct in the adult. Ho homologue in the male exactly. 

Paradidymis or organ of Giraldes in males and free-martins;- the remains 
of the caudal group of mesonephric tubules, homologous tc the 
paroophoron of tho female. In males and free-ma-tins a single 
isolated tubule forms the Ductus aberrans. 

(UcMurrick) 

Priory sex-cords;- the first set of tubule primordia that invacinate 
from the germinal epithelium either type of gonad, and cut off 
from this epithelium by the primary tunica albuginea. 

(Hc'iiurrick) 

Prostate glands;- socalled homologueo of the para-urethral ducts of Skeene 
found in females and free-rsartincj prostatic 

urethra and vestibule. 

( Neugebauer, -‘c Airrick ) 








f 

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3 


Prontatic urethra;- the sale hcmologue to th© vestibule or vectibulu® 
vaginae of the feaale and flpii iwrtillllttit port of the einus uro- 
genitalii. derivatives between the opor.ing of the vagina a»i th© 
openings of the bulbo-ur ,thral or bulbo-vostibular glands to it. 
(liougebauer, VtoMurrick) 

Pete teatie or rote owU^ thooa anter* r cell-c ■'?:ians that grow ft"#* 
the genital ridge epithelium into th© strora ofth i raosonerhros, 
and their derivatives, rets oorda and rote tubules, 

(dc&urrick) 

Saccus vaginalis in wales and free-curtins;- the derivative of the 
vaginal process of the peritoneum or th© canal o** ftuck. 

(Mcdurrick, 

Secondary sex-cords or Qords cf Pfiftger;- secondary proliferations of the 
germinal epitheiiun into the interior of th gonad after the pri¬ 
mary ones are cut of : by the primary tunica albtuinea, and later 
cut off from the ge cinai epithelium by a secondary tunica. Ho 
hooologue in the testis. 

(Ucteurrick.; 

Seminal veaicles in wales and free- artino; hcnolo,.ues of the partner’s 
ducts of females; on© or more glands with their ducts that develop 
as lat ral diverticula fres the uasa dsf.rontia or olf Ian ducts. 
(Prenti3B and Arey) 

Sub-urethral diverticulum females and soma free-mrti;s ? without hoiso- 
lopua in the males a pouch projecting forward under the urethra 
from the* point of entry of th vagina into th© vestibule. 

(Sisson) 

Supra-urethral diverticulum o'* Males oniy^with no hcmolo.ue iri females 

or free-nartins:- a snail pouch loading dorsolly fro® the aerabranous 
urethra poster or to the openings of the bulbo~urethral glands. 

Tunica albuginea. Primary;- the layers of connective tissue primordius 
that separate the primary ssx-cords fre® the epithelium o* the 
gonad coat, 

(Sillier) 







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4 . 


Tunica albuginea, iooondaryr- th'- definitive tunica of tho o*ary;- 

the lay-ra o*' connective tissue thut ep .r^te tl • aocoud&ry sax- 
cords free t ho epithelial cover cf the gcn&d in tho female. 

(WIliter) 

Tunica v&eoulore;- the vascular region inside the primary tunica al¬ 
buginea and between it and the prin&ry sx~cords. 

(Willier) 

Urethra proper;- or pars proxinmli© vosicao;- male homologue of tho 
urethra of the female and free-rartin;- that part of tho urethra 
between the bases of the; ureters and the ^fclffian ducts. 
(Keu&eb&uer) 

Uterus mueculinus;- da e hooologuo of tho Corpus uteri of feaules;- 
the derivatives of the part of tho fused !4$llerian ducts nearest 
the point of union of tho genital cords. Cornua uteri are those 
parte of the Mullerian duc f s uet anterior to the union of the 
genital cords and enlarged. 

Vagina masculine, Prostotic utriculus, or Sinus pocularis;- the male 
hossologu? of the vagina of the female;- the derivative of the 
enlarged posterior end of the united t&llerian duct©. 

(Prentiss and Arey) 

. \ 

Vulva of the female and fr®e-r?.rtin with no honologue in the male 5 - 
the perforated uroren tai plate and its derivatives,-used in the German 

C< y) 

sense as outlined by Sisson; Anat r.y cf the Domestic Animals. 

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Deacidified using the Bookkeeper process. 
Neutralizing agent: Magnesium Oxide 
Treatment Date: Dec. 2012 




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